{"id":1675,"date":"2026-08-20T17:13:50","date_gmt":"2026-08-20T21:13:50","guid":{"rendered":"https:\/\/opentextbooks.concordia.ca\/explorations3\/?post_type=chapter&#038;p=1675"},"modified":"2026-09-01T13:35:10","modified_gmt":"2026-09-01T17:35:10","slug":"1675","status":"publish","type":"chapter","link":"https:\/\/opentextbooks.concordia.ca\/explorations3\/chapter\/1675\/","title":{"raw":"Meet the Living Primates","rendered":"Meet the Living Primates"},"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 class=\"import-Normal\">Describe how studying nonhuman primates is important in anthropology.<\/li>\r\n \t<li class=\"import-Normal\">Compare two ways of categorizing taxa: grades and clades.<\/li>\r\n \t<li class=\"import-Normal\">Define different types of traits used to evaluate primate taxa.<\/li>\r\n \t<li class=\"import-Normal\">Identify key ways that primates differ from other mammals.<\/li>\r\n \t<li class=\"import-Normal\">Distinguish between the major primate taxa using their key characteristics.<\/li>\r\n \t<li class=\"import-Normal\">Describe your place in nature by learning your taxonomic classification.<\/li>\r\n<\/ul>\r\n<\/div>\r\n<\/div>\r\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>\r\n\r\n<h2 class=\"import-Normal\">Studying Primates in Biological Anthropology<\/h2>\r\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>[pb_glossary id=\"944\"]homology[\/pb_glossary], <\/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>\r\n<p class=\"import-Normal\">Nonhuman primates also make excellent comparators for learning about humans via <strong>[pb_glossary id=\"948\"]analogy[\/pb_glossary]<\/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>\r\n\r\n<h3 class=\"import-Normal\"><strong>Ways of Organizing Taxa<\/strong><\/h3>\r\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>[pb_glossary id=\"1116\"]clade[\/pb_glossary] <\/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>[pb_glossary id=\"1118\"]African clade[\/pb_glossary]<\/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>[pb_glossary id=\"1120\"]Asian clade[\/pb_glossary]<\/strong> of hominoids.<\/p>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"800\"]<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\" \/> 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>.][\/caption]\r\n<p class=\"import-Normal\">In contrast, <strong>[pb_glossary id=\"1122\"]grades[\/pb_glossary] <\/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>\r\n\r\n<h3 class=\"import-Normal\"><strong>Types of Traits<\/strong><\/h3>\r\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>[pb_glossary id=\"1124\"]Ancestral traits[\/pb_glossary]<\/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>[pb_glossary id=\"1126\"]Derived traits[\/pb_glossary]<\/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>\r\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>\r\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>[pb_glossary id=\"1128\"]Generalized traits[\/pb_glossary] <\/strong>are those characteristics that are useful for a wide range of things. Having <strong>[pb_glossary id=\"1138\"]opposable thumbs[\/pb_glossary]<\/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>[pb_glossary id=\"1130\"]Specialized traits[\/pb_glossary] <\/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>\r\n\r\n<h2 class=\"import-Normal\">What Makes Something a Primate?<\/h2>\r\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>\r\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>[pb_glossary id=\"1132\"]postorbital bar[\/pb_glossary], <\/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>[pb_glossary id=\"1134\"]postorbital plate[\/pb_glossary] <\/strong>or<strong> [pb_glossary id=\"1136\"]postorbital closure[\/pb_glossary] <\/strong>in addition to the bar (Figure 6.2).The postorbital bar is a derived trait of primates, appearing in our earliest ancestors.<\/p>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"661\"]<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\" \/> 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>.][\/caption]\r\n<p class=\"import-Normal\">Another distinctive trait of our Order is that many primates have <strong>[pb_glossary id=\"1140\"]trichromatic color vision[\/pb_glossary]<\/strong>, the ability to distinguish reds and yellows in addition to blues and greens. Birds, fish, and reptiles are <strong>[pb_glossary id=\"1142\"]tetrachromatic[\/pb_glossary] <\/strong>(they can see reds, yellows, blues, greens, and even ultraviolet), but most mammals, including some primates, are only <strong>[pb_glossary id=\"1144\"]dichromatic[\/pb_glossary] <\/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>\r\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>[pb_glossary id=\"1146\"]evolutionary trade-offs[\/pb_glossary]<\/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>\r\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>\r\n<p class=\"import-Normal\">Animals with large brains usually have extended life history patterns, and primates are no exception. <strong>[pb_glossary id=\"1148\"]Life history[\/pb_glossary] <\/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>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"392\"]<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\" \/> 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>.[\/caption]\r\n\r\nPrimates also differ from other animals in our hands and feet. The Order Primates is a largely <strong>[pb_glossary id=\"1150\"]arboreal[\/pb_glossary]<\/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>[pb_glossary id=\"1152\"]pentadactyly[\/pb_glossary]<\/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>[pb_glossary id=\"1154\"]terrestrial[\/pb_glossary]<\/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>[pb_glossary id=\"1156\"]tactile pads[\/pb_glossary] <\/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.\r\n<table class=\"alignright\" style=\"width: 219.75pt;height: 407px\"><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>\r\n<thead>\r\n<tr style=\"height: 30px\">\r\n<td class=\"a-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 361.667px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Primate suite of traits<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a-R\" style=\"height: 362px\">\r\n<td class=\"a-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 362px;width: 361.667px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Convergent eyes<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital bar<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Many have trichromatic colour vision<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short snouts<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Opposable thumbs and big toes<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Pentadactyly<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Flattened nails<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Tactile pads<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Highly arboreal<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Large brains<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Extended life histories<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Live in the tropics<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr style=\"height: 15px\">\r\n<td style=\"height: 15px;width: 362.033px\"><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\nLastly, 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.\r\n<h2 class=\"import-Normal\">Key Traits Used to Distinguish Between Primate Taxa<\/h2>\r\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>\r\n\r\n<h3 class=\"import-Normal\"><strong>Dental Characteristics<\/strong><\/h3>\r\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>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"356\"]<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\" \/> 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>.[\/caption]\r\n\r\nLike other mammals, primates are <strong>[pb_glossary id=\"1158\"]heterodont[\/pb_glossary]<\/strong>: they have multiple types of teeth that are used for different purposes. We have <strong>[pb_glossary id=\"1160\"]incisors[\/pb_glossary] <\/strong>for slicing; <strong>[pb_glossary id=\"1162\"]premolars[\/pb_glossary] <\/strong>and <strong>[pb_glossary id=\"1164\"]molars[\/pb_glossary] <\/strong>for grinding up our food; and <strong>[pb_glossary id=\"1166\"]canines[\/pb_glossary]<\/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>[pb_glossary id=\"1168\"]sexually dimorphic[\/pb_glossary]<\/strong>, with males tending to have larger canines than females. Some nonhuman primates <strong>[pb_glossary id=\"1170\"]hone[\/pb_glossary]<\/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>[pb_glossary id=\"1172\"]diastema[\/pb_glossary]<\/strong> (Figure 6.5).\r\n<p class=\"import-Normal\">We use a <strong>[pb_glossary id=\"1174\"]dental formula[\/pb_glossary]<\/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>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"241\"]<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\" \/> 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>.[\/caption]\r\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>[pb_glossary id=\"1176\"]cusps[\/pb_glossary]<\/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>\r\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>[pb_glossary id=\"954\"]frugivores[\/pb_glossary]<\/strong>), those who eat mostly insects (<strong>[pb_glossary id=\"958\"]insectivores[\/pb_glossary]<\/strong>), and those who eat primarily leaves (<strong>[pb_glossary id=\"956\"]folivores[\/pb_glossary]<\/strong>). A few primate taxa are <strong>[pb_glossary id=\"1178\"]gummivores[\/pb_glossary]<\/strong>, specializing in eating gums and saps, but we will only focus on the adaptations found in the three primary dietary groups.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Frugivores<\/em><\/h4>\r\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>[pb_glossary id=\"1180\"]bunodont[\/pb_glossary] <\/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>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"503\"]<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\" \/> 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>[\/caption]\r\n<h4 class=\"import-Normal\"><em>Insectivores<\/em><\/h4>\r\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>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"351\"]<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\" \/> 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>.][\/caption]\r\n<h4 class=\"import-Normal\"><em>Folivores<\/em><\/h4>\r\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>[pb_glossary id=\"1182\"]shearing crests[\/pb_glossary]<\/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>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"468\"]<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\" \/> 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>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Behavioural Adaptations<\/strong><\/h3>\r\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>[pb_glossary id=\"1184\"]activity patterns[\/pb_glossary]<\/strong>: whether they are active during the day (<strong>[pb_glossary id=\"1012\"]diurnal[\/pb_glossary]<\/strong>), at night (<strong>[pb_glossary id=\"1010\"]nocturnal[\/pb_glossary]<\/strong>), or through the 24-hour period (<strong>[pb_glossary id=\"1186\"]cathemeral[\/pb_glossary]<\/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>\r\n\r\n<h3 class=\"import-Normal\"><strong>Locomotor Adaptations<\/strong><\/h3>\r\n<p class=\"import-Normal\">Finally, primate groups vary in their adaptations for different forms of <strong>[pb_glossary id=\"1188\"]locomotion[\/pb_glossary]<\/strong>, or how they move around. Living primates are known to move by vertical clinging and leaping, quadrupedalism, brachiation, and bipedalism.<\/p>\r\n<p class=\"import-Normal\"><strong>[pb_glossary id=\"1190\"]Vertical clinging and leaping[\/pb_glossary] <\/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>\r\n\r\n<\/div>\r\n\r\n[caption id=\"attachment_181\" align=\"aligncenter\" width=\"608\"]<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\" \/> 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.)[\/caption]\r\n\r\n&nbsp;\r\n<div class=\"learning-objectives\">\r\n<p class=\"import-Normal\"><strong>[pb_glossary id=\"1192\"]Quadrupedalism[\/pb_glossary]<\/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>\r\n\r\n<\/div>\r\n\r\n[caption id=\"attachment_181\" align=\"aligncenter\" width=\"704\"]<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\" \/> 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>.[\/caption]\r\n\r\n<div class=\"learning-objectives\">\r\n<p class=\"import-Normal\">The third form of locomotion seen in primates is<strong> [pb_glossary id=\"1194\"]brachiation[\/pb_glossary]<\/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>[pb_glossary id=\"1196\"]semi-brachiation[\/pb_glossary]<\/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>[pb_glossary id=\"1198\"]prehensile tails[\/pb_glossary]<\/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>\r\n\r\n<\/div>\r\n\r\n[caption id=\"attachment_181\" align=\"alignnone\" width=\"1600\"]<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\" \/> 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>.[\/caption]\r\n\r\n<div class=\"learning-objectives\">\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"565\"]<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\" \/> 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>.[\/caption]\r\n<p class=\"import-Normal\">Lastly, humans move around on two feet, called <strong>[pb_glossary id=\"1200\"]bipedalism[\/pb_glossary]<\/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>\r\n\r\n<h2 class=\"import-Normal\">Primate Diversity<\/h2>\r\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>\r\n\r\n[caption id=\"\" align=\"alignnone\" width=\"2048\"]<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\" \/> 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>.][\/caption]\r\n<h3 class=\"import-Normal\"><strong>Suborder Strepsirrhini<\/strong><\/h3>\r\n[caption id=\"\" align=\"aligncenter\" width=\"387\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-1.jpg\" alt=\"Eight strepsirrhine species.\" width=\"387\" height=\"605\" \/> 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>.][\/caption]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>[pb_glossary id=\"1202\"]grooming claw[\/pb_glossary] <\/strong>(Figure 6.16) on the second digit of each foot, and the <strong>[pb_glossary id=\"1204\"]tooth comb[\/pb_glossary]<\/strong> (or <strong>[pb_glossary id=\"1206\"]dental comb[\/pb_glossary]<\/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.\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"237\"]<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\" \/> 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>[\/caption]\r\n<p class=\"import-Normal\">Compared to haplorrhines, strepsirrhines rely more on nonvisual senses. Strepsirrhines get their name because they have wet noses (<strong>[pb_glossary id=\"1208\"]rhinariums[\/pb_glossary]<\/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>[pb_glossary id=\"1210\"]scent marking[\/pb_glossary]<\/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>[pb_glossary id=\"1212\"]tapetum lucidum[\/pb_glossary]<\/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>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"292\"]<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\" \/> 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>[\/caption]\r\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>\r\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>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"443\"]<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\" \/> 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>.[\/caption]\r\n<h4><em>Lemurs of Madagascar<\/em><\/h4>\r\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>\r\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>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"326\"]<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\" \/> 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>.[\/caption]\r\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>\r\n\r\n<h4 class=\"import-Normal\"><em>Lorises, Pottos, and Galagos of Asia and Africa<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"207\"]<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\" \/> 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>.[\/caption]\r\n\r\nUnlike 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.\r\n\r\nThe 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.\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<thead>\r\n<tr>\r\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Lemurs<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Lorises, Pottos, and Galagos<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a0-R\">\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Geographic range<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Madagascar<\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">South and Southeast Asia<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Central Africa<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a0-R\">\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Activity patterns<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Diurnal, nocturnal, or cathemeral<\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Nocturnal<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a0-R\">\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Dietary types<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Insectivore, frugivore, or folivore<\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Insectivore, frugivore<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a0-R\">\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Social groupings<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary, pairs, or small to large groups<\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a0-R\">\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Forms of locomotion<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Vertical clinger leapers, quadrupedal<\/p>\r\n<\/td>\r\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Slow quadrupedal climbers and active quadrupedal runners<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<h3 class=\"import-Normal\"><strong>Suborder Haplorrhini<\/strong><\/h3>\r\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>\r\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>[pb_glossary id=\"1410\"]fovea[\/pb_glossary]<\/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>[pb_glossary id=\"1412\"]dry nose[\/pb_glossary] <\/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>\r\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>[pb_glossary id=\"1414\"]monomorphic[\/pb_glossary]<\/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>\r\n\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<thead>\r\n<tr>\r\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Suborder Haplorrhini<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a1-R\">\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Sensory adaptations<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Rhinarium<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Longer snout<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Eyes less convergent<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital bar<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Tapetum lucidum<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Mobile ears<\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">No rhinarium<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short snout<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Eyes more convergent<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital plate<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">No tapetum lucidum<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Many are trichromatic<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Fovea<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a1-R\">\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Dietary differences<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly insectivores and frugivores, few folivores<\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Few insectivores, mostly frugivores and folivores<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a1-R\">\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Activity patterns and Ecology<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly nocturnal, few diurnal or cathemeral<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Almost entirely arboreal<\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Only two are nocturnal, rest are diurnal<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Many arboreal taxa, also many terrestrial taxa<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a1-R\">\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Social groupings<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly solitary, some pairs, small to large groups<\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Only two are solitary, all others live in pairs, small to very large groups<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a1-R\">\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Sexual dimorphism<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Minimal to none<\/p>\r\n<\/td>\r\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Few taxa have little\/none, many taxa show moderate to high dimorphism<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\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>\r\n\r\n<h4 class=\"import-Normal\"><em>Infraorder Tarsiiformes of Asia<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"188\"]<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\" \/> 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>.[\/caption]\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"362\"]<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\" \/> 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>.[\/caption]\r\n\r\nToday, 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>[pb_glossary id=\"1416\"]faunivorous[\/pb_glossary] <\/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.\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"486\"]<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\" \/> 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>[\/caption]\r\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>\r\n\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<thead>\r\n<tr>\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Like Strepsirrhini<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Unique<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Like Haplorrhini<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a2-R\">\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Very small<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Nocturnal<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Highly insectivorous<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Vertical clinger-leapers<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Little\/no sexual dimorphism<\/p>\r\n<\/td>\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Two grooming claws<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:3:3\/1:1:3:3 dental formula<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Do not eat vegetation<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Can rotate their heads nearly 180 degrees<\/p>\r\n<\/td>\r\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Almost full PO closure<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">More convergent eyes<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">No tapetum lucidum<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">No rhinarium<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Genetic evidence<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Fovea<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<h4 class=\"import-Normal\"><em>Infraorder Platyrrhini of Central and South America<\/em><\/h4>\r\n[caption id=\"attachment_179\" align=\"alignleft\" width=\"329\"]<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\" \/> 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>.[\/caption]\r\n\r\nThe 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.\r\n\r\nOn 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>[pb_glossary id=\"1418\"]polymorphic color vision[\/pb_glossary]<\/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>[pb_glossary id=\"1420\"]monochromatic[\/pb_glossary]<\/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.\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"279\"]<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\" \/> 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>.[\/caption]\r\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>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"428\"]<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\" \/> 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>].][\/caption]\r\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>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"458\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image33-1.jpg\" alt=\"Four platyrrhine species.\" width=\"458\" height=\"457\" \/> 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>.][\/caption]\r\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>\r\n\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<tbody>\r\n<tr class=\"a3-R\">\r\n<td class=\"a3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Platyrrhini traits<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a3-R\">\r\n<td class=\"a3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Flat nose with rounded nostrils pointing to the side<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Highly arboreal<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Less sexually dimorphic on average<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:3:3 dental formula*<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Polymorphic colour vision*<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<h4 class=\"import-Normal\"><em>Infraorder Catarrhini of Asia and Africa <\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"191\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image34.jpg\" alt=\"Wolf\u2019s guenon.\" width=\"191\" height=\"287\" \/> 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>.[\/caption]\r\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>\r\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>\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"632\"]<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\" \/> 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>.][\/caption]\r\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>\r\n\r\n<h4 class=\"import-Normal\"><em>Superfamily Cercopithecoidea of Africa and Asia<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"285\"]<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\" \/> 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>[\/caption]\r\n\r\nCompared to hominoids, cercopithecoids have an ancestral quadrupedal body plan with two key derived traits. The first derived trait of cercopithecoids is their <strong>[pb_glossary id=\"1422\"]bilophodont[\/pb_glossary] <\/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>[pb_glossary id=\"1424\"]ischial callosities[\/pb_glossary] <\/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.\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"359\"]<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\" \/> 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>.[\/caption]\r\n\r\nCercopithecoid 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.\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"180\"]<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\" \/> 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>[\/caption]\r\n\r\nCercopithecoidea 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>[pb_glossary id=\"1426\"]natal coats[\/pb_glossary]<\/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.\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"408\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image39.jpg\" alt=\"Male proboscis monkey.\" width=\"408\" height=\"272\" \/> 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>.[\/caption]\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"414\"]<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\" \/> 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>.[\/caption]\r\n<h4 class=\"import-Normal\"><em>Superfamily Hominoidea of Africa and Asia<\/em><\/h4>\r\n[caption id=\"\" align=\"aligncenter\" width=\"438\"]<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\" \/> 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>.[\/caption]\r\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>[pb_glossary id=\"1428\"]Y-5 molars[\/pb_glossary]<\/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>\r\nTo 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>[pb_glossary id=\"1430\"]olecranon process[\/pb_glossary]<\/strong>, which allows for improved extension in our arms. At the wrist end of the ulna, hominoids have a short <strong>[pb_glossary id=\"1432\"]styloid process[\/pb_glossary]<\/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.\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<thead>\r\n<tr>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Quadrupedalism<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Brachiation<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Arm length vs. leg length<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">About equal<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Arms are longer<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Shoulder position<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">More on the front<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Out to the side<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ribcage shape<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Deep front-to-back<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Narrow side-to-side<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Shallow front-to-back<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Wide side-to-side<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Length of lower back<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Collar bone length<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ulnar olecranon process<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ulnar styloid process<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"a4-R\">\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Tail<\/strong><\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Short to long<\/p>\r\n<\/td>\r\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">None<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\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>\r\n\r\n<table class=\"aligncenter\"><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>\r\n<thead>\r\n<tr style=\"height: 22pt\">\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\" colspan=\"2\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Infraorder Catarrhini<\/strong><\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a5-R\" style=\"height: 22pt\">\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\" colspan=\"2\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Downward facing, tear-drop shaped nostrils, close together<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Arboreal and more terrestrial taxa<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">On average, largest primates<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">On average, most sexually dimorphic taxonomic group<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:2:3 dental formula<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">All trichromatic<\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<table class=\"aligncenter\" style=\"width: 468pt\"><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>\r\n<thead>\r\n<tr>\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<\/td>\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\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>\r\n<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"a5-R\">\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Wide geographic distribution<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Bilophodont molars<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Ischial callosities<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Reproduce every 1\u20132 years<\/p>\r\n<\/td>\r\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\">Tropical forests of Africa and Asia<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Y-5 molars<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Adaptations for brachiation<\/p>\r\n<p class=\"import-Normal\" style=\"text-align: center\">Reproduce every 4\u20139 years<\/p>\r\n<\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<h4 class=\"import-Normal\"><em>Family <\/em>Hylobatidae <em>of Southeast Asia<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"441\"]<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\" \/> 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>.[\/caption]\r\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>\r\n\r\n<h4 class=\"import-Normal\">Pongo<em> of Southeast Asia<\/em><\/h4>\r\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>[pb_glossary id=\"1434\"]sagittal crest[\/pb_glossary]<\/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>[pb_glossary id=\"1436\"]male bimaturism[\/pb_glossary]<\/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>\r\n&nbsp;\r\n\r\n<\/div>\r\n\r\n[caption id=\"attachment_181\" align=\"aligncenter\" width=\"1900\"]<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\" \/> 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>.[\/caption]\r\n\r\n<div class=\"learning-objectives\">\r\n<h4 class=\"import-Normal\">Gorilla <em>of Africa<\/em><\/h4>\r\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>[pb_glossary id=\"1438\"]knuckle-walking[\/pb_glossary]<\/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>\r\n\r\n<\/div>\r\n\r\n[caption id=\"attachment_181\" align=\"aligncenter\" width=\"1900\"]<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\" \/> 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>.[\/caption]\r\n\r\n<div class=\"learning-objectives\">\r\n<h4 class=\"import-Normal\">Pan<em> of Africa<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"252\"]<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\" \/> 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>.[\/caption]\r\n\r\nThe 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.\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"418\"]<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\" \/> 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>.[\/caption]\r\n<h4 class=\"import-Normal\">Homo<\/h4>\r\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>\r\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>\r\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>\r\n\r\n<div class=\"textbox\">\r\n<h2 class=\"import-Normal\">Special Topic: Primates in Culture and Religion<\/h2>\r\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>\r\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>\r\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>\r\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>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"308\"]<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\" \/> 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>.[\/caption]\r\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>\r\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>\r\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>[pb_glossary id=\"1440\"]Ethnoprimatology[\/pb_glossary]<\/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>\r\n\r\n<\/div>\r\n<h2 class=\"import-Normal\">Summary<\/h2>\r\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>\r\n\r\n<\/div>\r\n<h2>Expand your view<\/h2>\r\nFor 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>\r\n\r\n&nbsp;\r\n<div class=\"learning-objectives\">\r\n<div class=\"textbox shaded\">\r\n<h2 class=\"import-Normal\">Review Questions<\/h2>\r\n<ul>\r\n \t<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>\r\n \t<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>\r\n \t<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>\r\n \t<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>\r\n \t<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>\r\n<\/ul>\r\n<\/div>\r\n<h2 class=\"import-Normal\">Key Terms<\/h2>\r\n<p class=\"import-Normal\"><strong>Activity pattern<\/strong>: Refers to the time of day an animal is typically active.<\/p>\r\n<p class=\"import-Normal\"><strong>African clade<\/strong>: A grouping that includes gorillas, chimpanzees, bonobos, humans, and their extinct relatives.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Ancestral trait<\/strong>: A trait that has been inherited from a distant ancestor.<\/p>\r\n<p class=\"import-Normal\"><strong>Arboreal<\/strong>: A descriptor for an organism that spends most of its time in trees.<\/p>\r\n<p class=\"import-Normal\"><strong>Asian clade<\/strong>: A grouping that includes orangutang and their extinct relatives.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Bipedalism<\/strong>: Walking on two legs.<\/p>\r\n<p class=\"import-Normal\"><strong>Brachiation<\/strong>: A form of locomotion in which the organism swings below branches using the forelimbs.<\/p>\r\n<p class=\"import-Normal\"><strong>Bunodont<\/strong>: Low, rounded cusps on the cheek teeth.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Cathemeral<\/strong>: Active throughout the 24-hour period.<\/p>\r\n<p class=\"import-Normal\"><strong>Clade<\/strong>: A grouping based on ancestral relationships; a branch of the evolutionary tree.<\/p>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Derived trait<\/strong>: A trait that has been recently modified, most helpful when assigning taxonomic classification.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Dichromatic<\/strong>: Being able to see only blues and greens.<\/p>\r\n<p class=\"import-Normal\"><strong>Diurnal<\/strong>: Active during the day.<\/p>\r\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>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Faunivorous<\/strong>: Having a diet consisting entirely of animal matter: insects, eggs, lizards, etc.<\/p>\r\n<p class=\"import-Normal\"><strong>Folivore<\/strong>: Having a diet consisting primarily of leaves.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Frugivore<\/strong>: Having a diet consisting primarily of fruit.<\/p>\r\n<p class=\"import-Normal\"><strong>Generalized trait<\/strong>: A trait that is useful for a wide range of tasks.<\/p>\r\n<p class=\"import-Normal\"><strong>Grade<\/strong>: A grouping based on overall similarity in lifestyle, appearance, and behaviour.<\/p>\r\n<p class=\"import-Normal\"><strong>Grooming claw<\/strong>: A claw present on the second pedal digit in strepsirrhines.<\/p>\r\n<p class=\"import-Normal\"><strong>Gummivore<\/strong>: Having a diet consisting primarily of gums and saps.<\/p>\r\n<p class=\"import-Normal\"><strong>Heterodont<\/strong>: Having different types of teeth.<\/p>\r\n<p class=\"import-Normal\"><strong>Homology<\/strong>: When two or more taxa share characteristics because they inherited them from a common ancestor.<\/p>\r\n<p class=\"import-Normal\"><strong>Hone<\/strong>: When primates sharpen their canines by wearing them on adjacent teeth.<\/p>\r\n<p class=\"import-Normal\"><strong>Incisors<\/strong>: The spatula-shaped teeth at the front of the mouth.<\/p>\r\n<p class=\"import-Normal\"><strong>Insectivore<\/strong>: Having a diet consisting primarily of insects.<\/p>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Life history<\/strong>: Refers to an organism\u2019s pace of growth, reproduction, lifespan, etc.<\/p>\r\n<p class=\"import-Normal\"><strong>Locomotion<\/strong>: How an organism moves around.<\/p>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Monochromatic<\/strong>: Being able to see only in shades of light to dark, no colour.<\/p>\r\n<p class=\"import-Normal\"><strong>Monomorphic<\/strong>: When males and females of a species do not exhibit significant sexual dimorphism.<\/p>\r\n<p class=\"import-Normal\"><strong>Natal coat<\/strong>: Refers to the contrasting fur colour of baby leaf monkeys compared to adults.<\/p>\r\n<p class=\"import-Normal\"><strong>Nocturnal<\/strong>: Active at night.<\/p>\r\n<p class=\"import-Normal\"><strong>Olecranon process<\/strong>: Bony projection at the elbow end of the ulna.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Pentadactyly<\/strong>: Having five digits or fingers and toes.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Postorbital bar<\/strong>: A bony ring that surrounds the eye socket, open at the back.<\/p>\r\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>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Quadrupedalism<\/strong>: Moving around on all fours.<\/p>\r\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>\r\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>\r\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>\r\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>\r\n<p class=\"import-Normal\"><strong>Sexually dimorphic<\/strong>: When a species exhibits sex differences in morphology, behaviour, hormones, and\/or colouration.<\/p>\r\n<p class=\"import-Normal\"><strong>Shearing crests<\/strong>: Sharpened ridges that connect cusps on a bilophodont molar.<\/p>\r\n<p class=\"import-Normal\"><strong>Specialized trait<\/strong>: A trait that has been modified for a specific purpose.<\/p>\r\n<p class=\"import-Normal\"><strong>Styloid process of ulna<\/strong>: A bony projection of the ulna at the end near the wrist.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Tapetum lucidum<\/strong>: Reflecting layer at the back of the eye that magnifies light.<\/p>\r\n<p class=\"import-Normal\"><strong>Terrestrial<\/strong>: A descriptor for an organism that spends most of its time on the ground.<\/p>\r\n<p class=\"import-Normal\"><strong>Tetrachromatic<\/strong>: Having the ability to see reds, yellows, blues, greens, and ultraviolet.<\/p>\r\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>\r\n<p class=\"import-Normal\"><strong>Trichromatic colour vision<\/strong>: Being able to distinguish yellows and reds in addition to blues and greens.<\/p>\r\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>\r\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>\r\n\r\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\r\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>\r\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>\r\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution<\/em>. Third edition. San Diego: Academic Press.<\/p>\r\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>\r\n<p class=\"import-Normal\">Rowe, Noel. 1996. <em>Pictorial Guide to the Living Primates<\/em>. Charlestown, RI: Pogonias Press.<\/p>\r\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>\r\n\r\n<h2>References<\/h2>\r\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>\r\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>\r\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>\r\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>\r\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>\r\n<p class=\"import-Normal\">Dunbar, Robin I. M. 1998. \u201cThe Social Brain Hypothesis.\u201d <em>Evolutionary Anthropology<\/em> 6 (5): 178\u2013190.<\/p>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\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>\r\n\r\n<h2>Acknowledgements<\/h2>\r\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>\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 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_1675_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_1675_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_1675_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_1675_1120\">Asian clade<\/a><\/strong> of hominoids.<\/p>\n<figure style=\"width: 800px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_1134\">postorbital plate<\/a> <\/strong>or<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 &#8230;. (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_1675_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_1675_1160\">incisors<\/a> <\/strong>for slicing; <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_1162\">premolars<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_1164\">molars<\/a> <\/strong>for grinding up our food; and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_956\">folivores<\/a><\/strong>). A few primate taxa are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_1012\">diurnal<\/a><\/strong>), at night (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_1010\">nocturnal<\/a><\/strong>), or through the 24-hour period (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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&#8217;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_1675_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_1675_1204\">tooth comb<\/a><\/strong> (or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 &#8211; 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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&#8217;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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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_1675_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 &#8211; 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 &#8211; 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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_1675_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 loading=\"lazy\" decoding=\"async\" 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_1675_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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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 loading=\"lazy\" decoding=\"async\" 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_1675_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 class=\"glossary\"><span class=\"screen-reader-text\" id=\"definition\">definition<\/span><template id=\"term_1675_944\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_944\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sarah S. King, Ph.D., Cerro Coso Community College<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Kara Jones, M.A., Ph.D. student, University of Nevada Las Vegas<\/p>\n<h6>Student conbtributors for this chapter: Catherine Belec, Maria Papadakis, Camille Senior and Nadjat Baril<\/h6>\n<p class=\"import-Normal\"><em>This chapter<\/em><em> is a revision from \"<\/em><a class=\"rId6\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\"><em>Chapter 7: Understanding the Fossil Context<\/em><\/a><em>\u201d by Sarah King and Lee Anne Zajicek. <\/em><em>In <\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId8\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">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 analyze, and their applications.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Describe the methods used to reconstruct past environments.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Interpret a site using the methods described in this chapter.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fossil Study: An Evolving Process<\/h2>\n<h3 class=\"import-Normal\"><strong>Mary Anning and the Age of Wonder<\/strong><\/h3>\n<figure style=\"width: 206px\" class=\"wp-caption alignleft\"><img src=\"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-meter <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_1675_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_1675_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_1675_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 Center, China Lake, California, in 1945. The fossil was buried deep in the strata and was pulled out of the ground along with a crashed \u201cFat Boy\u201d missile after atomic missile testing (S. Brubaker, personal communication, March 9, 2018). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Murexsul (Figure 7.6)<\/a> from the <a href=\"https:\/\/maturango.org\/\">Maturango Museum<\/a>, Ridgecrest, California, by Sarah S. King and Lee Anne Zajicek is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>The work of Anning, Darwin, Lyell, and many others laid the foundation for the modern methods we use today. Though anthropology is focused on humans and our primate relatives (and not on dinosaurs, as many people wrongly assume), you will see that methods developed in paleontology, geology, chemistry, biology, and physics are often applied in anthropological research. In this chapter, you will learn about the primary methods and techniques employed by biological anthropologists to answer questions about <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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 analyze evidence and reconstruct past activities and environments.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Earth: It's Older than Dirt<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientists have developed precise and accurate dating methods based on work in the fields of physics and chemistry. Using these methods, scientists are able to establish the age of Earth as well as approximate ages of the organisms that have lived here. Earth is roughly 4.6 billion years old, give or take a few hundred million years. The first evidence for a living organism appeared around 3.5 billion years ago (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_836\">eras<\/a>,<\/strong> and eras are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_838\">periods<\/a><\/strong>. Finally, periods are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_840\">Holocene<\/a><\/strong>. It began almost twelve thousand years ago (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_860\">petrified wood<\/a><\/strong>. This fossil is created when actual pieces of wood\u2014such as the trunk of a tree\u2014mineralize and turn into rock. Petrified wood is a combination of silica, calcite, and quartz, and it is both heavy and brittle. Petrified wood can be colorful and is generally aesthetically pleasing because all the features of the original tree\u2019s composition are illuminated through mineralization (Figure 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_1675_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_1675_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 color. 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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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-meter trail of hominin footprints were discovered at the same site. These 70 footprints, now referred to as the Laetoli Footprints, were created when early humans walked in wet volcanic ash. Before the impressions were obscured, more volcanic ash and rain fell, sealing the footprints. These series of environmental events were truly extraordinary, but they fortunately resulted in some of the most famous and revealing trace fossils ever found. Dating of the footprints indicate that they were made 3.6 mya (Smithsonian National Museum of Natural History 2018).<\/p>\n<figure style=\"width: 495px\" class=\"wp-caption alignleft\"><img src=\"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 center their values when conducting work with their ancestors. A good source for considering places to look for archaeological research ethics before conducting fieldwork (and ideally during your research design) is the 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 analyze the evolution of a fossil species or artifacts. To answer the question \u201cHow do we know what we know?,\u201d you have to know how archaeologists and paleoanthropologists establish dates for artifacts, fossils, and sites.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Though accurate dating is important for context and analysis, we must consider the impact. Many of the chronometric dating methods used by anthropologists require the removal of small samples from artifacts, bones, soils, and rock. Thus these techniques are considered destructive. How much of an artifact are you willing to destroy to get your date? Sharon Clough, a Senior Environmental Officer at Cotswold Archaeology, addressed this issue in a case study from her research. She stated that \u201cthe benefit of a date did not outweigh the destruction of a valuable and finite resource\u201d (Clough 2020). The resource in question was human remains. When considering our dating options, we want to be sure that we do as little harm as possible, especially in the case of human remains (read more about this issue in the Special Topic box, \u201cNecropolitics\u201d).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dating techniques are divided into two broad categories: relative dating methods and chronometric (sometimes called absolute) dating methods.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_890\">Elements<\/a>.<\/strong>\u00a0Each element consists of an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_904\">dendrochronology<\/a><\/strong> below), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_908\">Potassium-argon (K-Ar) dating<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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.\u00a0Another 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_1675_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_1675_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_1675_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_1675_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 catalog of tree profiles (see University of Arizona n.d.). Archaeologists can then compare wood artifacts and ecofacts with existing timelines, provided the tree rings are visible, and find where their artifacts fit in the pattern. Dendrochronology has been in use since the early 20th century (Dean 2009, 25). The Northern Hemisphere chronology stretches back nearly 14,000 years (Reimer Et al. 2013, 1870) and has been used successfully to date southwestern U.S. sites such as Pueblo Bonito and Aztec Ruin (Dean 2009, 26). Dendrochronological evidence has helped calibrate radiocarbon dates and even provided direct evidence of global warming (Dean 2009, 26\u201327).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In Australia, dendrochronology, along with other environmental reconstruction methods, has been used to show that the Indigenous people had sophisticated land management systems before the arrival of British invaders. According to the work of Michael-Shawn Fletcher and colleagues (2021), there was a significant encroachment of the rainforests and tree species into grasslands after the British invasion. Prior to this time, Indigenous people managed the landscape through controlled burns at regular intervals. This practice created climate-resistant grasslands that were biodiverse and provided predictable food supplies for humans and other animals. Under European land management, there have been negative impacts on biodiversity and climate resilience and an increase in catastrophic wildfires (Fletcher Et al. 2021). 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 analyze <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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 analyze the bones and teeth of an animal to identify the <sup>13<\/sup>C\/<sup>12<\/sup>C ratios and identify the types of plants that animal was eating.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The ratios of stable nitrogen isotopes <sup>15<\/sup>N and <sup>14<\/sup>N can also give information about the diet of fossilized or deceased organisms. Though initially absorbed from water and soils by plants, the nitrogen ratios change depending on the primary diet of the organism. An animal who has a mostly vegetarian diet will have lower ratios of <sup>15<\/sup>N to <sup>14<\/sup>N, while those further up the food chain, like carnivores, will have higher ratios of <sup>15<\/sup>N. Interestingly, breastfeeding infants have a higher nitrogen ratio than their mothers, because they are getting all of their nutrients through their mother\u2019s milk. So nitrogen can be used to track life events like weaning (Jay Et al. 2008, 2). A marine versus terrestrial diet will also affect the nitrogen signatures. Terrestrial diets have lower ratios of <sup>15<\/sup>N than marine diets. In the course of human evolution, this type of analysis can help us identify important changes in human nutrition. It can help anthropologists figure out when meat became a primary part of the ancient human diet or when marine resources began to be used. The ratios of stable nitrogen isotopes can also be used to determine a change in status, as in the case of the Llullaillaco children (the \u201cice mummies\u201d) found in the Andes Mountains. For instance, the nitrogen values in hair from the Llullaillaco Maiden showed a significant positive shift that is associated with increased meat consumption in the last 12 months of her life (Wilson Et al. 2007). Although the two younger children had little changes in their diets in the last year of their short lives, the changes in their nitrogen values were significant enough to suggest that the improvement in their diets may have been attributed to the Incas\u2019 desire to sacrifice healthy, high-status children\u201d (Faux 2012, 6).<\/p>\n<h4 class=\"import-Normal\"><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 analyzed, these isotopes would resemble those found in the geographic area where an individual lived as a child. Bones, however, are a different story. Bones are constantly changing throughout life. Old cells are removed and new cells are deposited to respond to growth, healing, activity change, and general deterioration. Therefore, the isotopic signature of bones will reflect the geographical area in which an individual spent the last seven to ten years of life. If an individual has different isotopic signatures for their bones and teeth, it could indicate a migration some time during their life after childhood.<\/p>\n<figure style=\"width: 386px\" class=\"wp-caption alignright\"><img src=\"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 analyzed in cranial fragments. Using the remains of 25 individuals, they compared their strontium signatures to the geology of Wessex and other regions of the UK. Fifteen of those individuals had strontium signatures that matched the local geology. This means that in the last ten or so years of their lives, they lived and ate food from around Stonehenge. However, ten of the individuals did not match the local geologic signature. These individuals had strontium ratios more closely aligned with the geology of west Wales. Archaeologists find this particularly interesting because in the early phases of Stonehenge\u2019s construction, the smaller \u201cblue stones\u201d were brought 200 km from Wales in a feat of early engineering. These larger regional connections show that Stonehenge was not just a site of local importance. It dominated a much larger region of influence and drew people from all over ancient Britain (Snoeck Et al. 2018).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Cold Case Naia<\/h2>\n<figure style=\"width: 455px\" class=\"wp-caption alignleft\"><img src=\"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_1675_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 analyze fossils to determine age, origins, and migration patterns as well as to provide insight into the health and diet of the fossilized organism. While each method has its advantages, disadvantages, and limited applications, these tools enable anthropologists to theorize how all living organisms evolved, including the evolution of early humans into modern humans, <em>H. sapiens<\/em>. The fossil record is far from complete, but our expanding understanding of the fossil context, with exciting new discoveries and improved scientific methods, enables us to document the history of our planet and the evolution of life on Earth.<\/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 analyzes the absorption of fluorine in bones from the surrounding soils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Foraminifera<\/strong>: Single-celled marine organisms with shells.<\/p>\n<p class=\"import-Normal\"><strong>Fossilization<\/strong>: The process by which an organism becomes a fossil.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Fossils<\/strong>: Mineralized copies of organisms or activity imprints.<\/p>\n<p class=\"import-Normal\"><strong>G<\/strong><strong>eomorphology<\/strong>: The study of the physical characteristics of the Earth\u2019s surface.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Glacial periods<\/strong>: Periods characterized by low global temperatures and the expansion of ice sheets on Earth\u2019s surface.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Holocene<\/strong>: The geologic epoch from 10 kya to present. (See the discussion on \u201cthe Anthropocene\u201d for the debate regarding the current epoch name.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Hominin<\/strong>: The term used for humans and their ancestors after the split with chimpanzees and bonobos.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>In matrix<\/strong>: When a fossil is embedded in a substance, such as igneous rock.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Isotopes<\/strong>: Variants of elements.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Kya<\/strong>: Thousand years ago.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Law of Superposition<\/strong>: The scientific law that states that rock and soil are deposited in layers, with the youngest layers on top and the oldest layers on the bottom.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Lithification<\/strong>: The process by which the pressure of sediments squeeze extra water out of decaying remains and replace the voids that appear with minerals from the surrounding soil and groundwater.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Luminescence dating<\/strong>: The chronometric dating method based on the buildup of background radiation in pottery, clay, and soils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Megafauna<\/strong>: Large animals such as mammoths and mastodons.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Mitochondrial DNA<\/strong>: DNA located in the mitochondria of a cell that is only passed down from biological mother to child.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Mya<\/strong>: Million years ago.<\/p>\n<p class=\"import-Normal\"><strong>P<\/strong><strong>aleopathology<\/strong>: Study of ancient diseases and injuries identified through examining remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Periods<\/strong>: Geologic time units that span millions of years and are subdivided into <em>epochs<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Permineralization<\/strong>: When minerals from water impregnate or replace organic remains, leaving a fossilized copy of the organism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Petrified wood<\/strong>: A fossilized piece of wood in which the original organism is completely replaced by minerals through petrifaction.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Potassium-argon (K-Ar) dating<\/strong>: A chronometric dating method that measures the ratio of argon gas in volcanic rock to estimate time elapsed since the volcanic rock cooled and solidified. See also <em>argon-argon dating<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Pseudofossils<\/strong>: Natural rocks or mineral formations that can be mistaken for fossils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Radioactive decay<\/strong>: The process of transforming the atom by spontaneously releasing energy.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Radiocarbon dating<\/strong>: The chronometric dating method based on the radioactive decay of <sup>14<\/sup>C in organic remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Relative dating<\/strong>: Dating methods that do not result in numbers of years but, rather, in relative timelines wherein some organisms or artifacts are older or younger than others.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Sediment cores<\/strong>: Core samples taken from lake beds or other water sources for analysis of their pollen.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Stable isotopes<\/strong>: Variants of elements that do not change over time without outside interference.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Stratigraphy<\/strong>: A relative dating method that is based on ordered layers or (strata) that build up over time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Taphonomy<\/strong>: The study of what happens to an organism after death.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Trace fossils<\/strong>: Fossilized remains of activity such as footprints.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Uniformitarianism<\/strong>: The theoretical perspective that the geologic processes observed today are the same as the processes operating in the past.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Unstable isotopes<\/strong>: Variants of elements that spontaneously change into stable isotopes over time.<\/p>\n<p class=\"import-Normal\"><strong>Uranium series dating<\/strong>: A radiometric dating method based on the decay chain of unstable isotopes of <sup>238<\/sup>U and <sup>235<\/sup>U.<\/p>\n<\/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. 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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\">Acknowledgments<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We are grateful to Lee Anne Zajicek, who coauthored the first edition. Her original contributions continue to be an integral part of this chapter. We thank the staff of the Maturango Museum, Ridgecrest, California. Specifically, for their generous help with photography and fossil images, we acknowledge Debbie Benson, executive director; Alexander K. Rogers, former archaeology curator; Sherry Brubaker, natural history curator; and Elaine Wiley, history curator. We thank Sharlene Paxton, a librarian at Cerro Coso Community College, Ridgecrest, California, for her guidance and expertise with OER and open-source images, and John Stenger-Smith and Claudia Sellers from Cerro Coso Community College, Ridgecrest, California, for their feedback on the chemistry and plant biology content. Finally, we thank William Zajicek and Lauren Zajicek, our community college students, for providing their impressions and extensive feedback on early drafts of the chapter.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_948\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_948\"><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_1675_1116\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1116\"><div tabindex=\"-1\"><p>The study of what happens to an organism after death.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1118\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1118\"><div tabindex=\"-1\"><p>An oxygen-free environment.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1120\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1120\"><div tabindex=\"-1\"><p>When minerals from water impregnate or replace organic remains, leaving a fossilized copy of the organism.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1122\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1122\"><div tabindex=\"-1\"><p>Million years ago.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1124\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1124\"><div tabindex=\"-1\"><p>Fossilized remains of activity such as footprints.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1126\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1126\"><div tabindex=\"-1\"><p>Fossilized poop.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1128\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1128\"><div tabindex=\"-1\"><p>Natural rocks or mineral formations that can be mistaken for fossils.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1138\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1138\"><div tabindex=\"-1\"><p>Variants of elements.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1130\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1130\"><div tabindex=\"-1\"><p>A relative dating method that is based on ordered layers or (strata) that build up over time.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1132\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1132\"><div tabindex=\"-1\"><p>A relative dating method that uses other plant and animal remains occurring in the stratigraphic context to establish time depth.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1134\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1134\"><div tabindex=\"-1\"><p>A relative dating method that analyzes the absorption of fluorine in bones from the surrounding soils.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1136\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1136\"><div tabindex=\"-1\"><p>Matter that cannot be broken down into smaller matter.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1140\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1140\"><div tabindex=\"-1\"><p>Variants of elements that spontaneously change into stable isotopes over time.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1142\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1142\"><div tabindex=\"-1\"><p>The chronometric dating method based on the radioactive decay of 14C in organic remains.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1144\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1144\"><div tabindex=\"-1\"><p>Single-celled marine organisms with shells.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1146\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1146\"><div tabindex=\"-1\"><p>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.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1148\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1148\"><div tabindex=\"-1\"><p>A chronometric dating method that is based on the fission of 283U.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1150\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1150\"><div tabindex=\"-1\"><p>A chronometric dating method that measures the background radiation accumulated in material over time.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1152\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1152\"><div tabindex=\"-1\"><p>The study of the physical characteristics of the Earth\u2019s surface.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1154\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1154\"><div tabindex=\"-1\"><p>DNA located in the mitochondria of a cell that is only passed down from biological mother to child.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1156\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1156\"><div tabindex=\"-1\"><p>The idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1158\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1158\"><div tabindex=\"-1\"><p>The scientific field that studies primate behavior and\/or ecology.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1160\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1160\"><div tabindex=\"-1\"><p>A trait with a function.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1162\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1162\"><div tabindex=\"-1\"><p>Understanding and highlighting the theory and research of non-Western individuals and perspectives.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1164\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1164\"><div tabindex=\"-1\"><p>A species that comes after the ancestor species.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1166\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1166\"><div tabindex=\"-1\"><p>The relationship between organisms and their physical surroundings.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1168\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1168\"><div tabindex=\"-1\"><p>Organisms whose diet consists primarily of fruit.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1170\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1170\"><div tabindex=\"-1\"><p>Organisms whose diets consist primarily of insects.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1172\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_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_1675_1174\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1174\"><div tabindex=\"-1\"><p>The branch of ecology that deals with the relationships and interactions between different organisms that occupy the same habitat.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1176\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1176\"><div tabindex=\"-1\"><p>Members of different species.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_954\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_954\"><div tabindex=\"-1\"><h2><img class=\"alignnone wp-image-584\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Katie-Nelson-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"250\" \/><\/h2>\n<h2>Katie Nelson, Ph.D.<\/h2>\n<h3>(Writer Chapter 1 &amp; 2, Editor Chapter 1-17)<\/h3>\n<p>Inver Hills Community College, kanelson@inverhills.edu<\/p>\n<p>Katie Nelson is an instructor of anthropology and sociology at Inver Hills Community College. She is the recipient of the 2022 Minnesota State Board of Trustees Educator of the Year award. Her research focuses on migration, identity, belonging, and citizenship(s) in human history and in the contemporary United States, Mexico, and Morocco.<\/p>\n<p>She received her B.A. in anthropology and Latin American studies from Macalester College, her M.A. in anthropology from the University of California, Santa Barbara, an M.A. in education and instructional technology from the University of Saint Thomas, and her Ph.D. from <a href=\"https:\/\/occidente.ciesas.edu.mx\/\">CIESAS Occidente (Centro de Investigaciones y Estudios Superiores en Antropologi\u0301a Socia<\/a>l \u2013Center for Research and Higher Education in Social Anthropology), based in Guadalajara, Mexico.<\/p>\n<p>Katie views teaching and learning as central to her practice as an anthropologist and is co-founder and Associate Editor of <a href=\"https:\/\/teachinglearninganthro.com\/\">Teaching and Learning Anthropology Journal<\/a>. She has contributed to several open access textbook projects, both as an author and an editor, and views the affordability of quality learning materials as an important piece of the equity and inclusion puzzle in higher education.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-585\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Lara-Braff-Author-Profile.jpg\" alt=\"\" width=\"218\" height=\"282\" \/><\/p>\n<h2>Lara Braff, Ph.D.<\/h2>\n<h3>(Writer Chapter 1 &amp; 2, Editor Chapter 1-17)<\/h3>\n<p>Grossmont College, Lara.Braff@gcccd.edu<\/p>\n<p>Lara Braff is a professor of anthropology at Grossmont College, where she teaches courses in cultural and biological anthropology. She received her B.A. in anthropology and Spanish from the University of California at Berkeley, and her M.A. and Ph.D. in comparative human development from the University of Chicago, where she specialized in medical anthropology.<\/p>\n<p>Lara\u2019s research, teaching, and involvement in open access projects (like this textbook) are rooted in concerns about social equity. In an effort to make college more accessible to all students, she serves as an Open Educational Resources (OER) coordinator at Grossmont College and Liaison for the Academic Senate for California Community Colleges\u2014Open Educational Resources Initiative.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-583\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Beth-Shook-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"300\" \/><\/p>\n<h2>Beth Shook, Ph.D.<\/h2>\n<h3>(Writer Chapter 1 &amp; 2, Editor Chapter 1-17)<\/h3>\n<p>California State University, Chico, bashook@csuchico.edu<\/p>\n<p>Beth Shook is a lecturer in the anthropology department at California State University, Chico. She received her B.A. in anthropology and in molecular biology from Cornell College (in Mount Vernon, Iowa) and her M.A. and Ph.D. in anthropology from the University of California, Davis. While she is broadly trained in anthropology, her research has focused on utilizing DNA in forensic and anthropological contexts.<\/p>\n<p>Beth enjoys teaching a variety of anthropology courses and mentoring graduate students in teaching. Additionally, she leads Chico State\u2019s Affordable Learning Solutions (CAL$) program, is committed to programs that prioritize diversity, and serves on the Society for Anthropology in Community Colleges (SACC) Executive Board.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-611\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Kelsie-Aguilera-Author-Profile-Small.jpeg\" alt=\"\" width=\"200\" height=\"200\" \/><\/p>\n<h2>Kelsie Aguilera, M.A.<\/h2>\n<h3>(Writer Chapter 1 &amp; 2, Editor Chapter 1-17)<\/h3>\n<p>Leeward Community College, kelsieag@hawaii.edu<\/p>\n<p>Kelsie Aguilera is an associate professor of anthropology at Leeward Community College. Located on the island of O\u2019ahu, Leeward Community College is part of the University of Hawai\u2019i System and holds a special commitment to Native Hawaiian education. At Leeward, Kelsie teaches anthropology courses in all of the subdisciplines.<\/p>\n<p>Kelsie received her B.A. in anthropology from the University of Miami and her M.A. in anthropology from Binghamton University. She is active within the American Anthropological Association and the Society for Anthropology in Community Colleges. She continues to work hard toward making anthropology accessible and relevant for her students.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-587\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Joylin-Namie-Author-Profile.png\" alt=\"\" width=\"200\" height=\"266\" \/><\/p>\n<h2>Joylin Namie, Ph.D.<\/h2>\n<h3>(Writer Chapter 3)<\/h3>\n<p>Truckee Meadows Community College, jnamie@tmcc.edu<\/p>\n<p>Joylin Namie is Professor of Anthropology at Truckee Meadows Community College, where she teaches courses in biological and cultural anthropology. Her current research interest is in (un)sustainable tourism in desert environments, particularly in the country of Jordan and the U.S. state of Nevada. She was awarded a fellowship to Jordan from the Council of American Overseas Research Centers (CAORC) in 2020 to explore this topic, including visiting Petra and other important tourism destinations in Jordan. Dr. Namie\u2019s favorite things in life are teaching, traveling, and spending time with her dog, Charley.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-588\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Hayley-Mann-Author-Profile.png\" alt=\"\" width=\"200\" height=\"259\" \/><\/p>\n<h2>Hayley Mann, M.A.<\/h2>\n<h3>(Writer Chapter 4)<\/h3>\n<p>Binghamton University, hmann3@binghamton.edu<\/p>\n<p>Hayley Mann received her bachelor\u2019s degree in Genetics from the University of California, Davis, and continued her graduate studies in Biological and Molecular Anthropology at the California State University, Sacramento. She is currently a Ph.D. candidate at Binghamton University, where her dissertation focus is on studying genetic variation of Pacific Islanders (Republic of Vanuatu) and also changes in health as the result of colonization. Hayley also works in clinical molecular carrier screening and specializes in DNA-sequencing procedures.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-612\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Andrea-J.-Alveshere-Author-Profile-Large.jpeg\" alt=\"\" width=\"200\" height=\"200\" \/><\/p>\n<h2>Andrea J. Alveshere, Ph.D.<\/h2>\n<h3>(Writer Chapter 5)<\/h3>\n<p>Western Illinois University, a-alveshere@wiu.edu, <a href=\"https:\/\/alveshere.com\/\">Andrea Alveshere: Archaeologist &amp; Biological Anthropologist<\/a><\/p>\n<p>Dr. Andrea Alveshere is an associate professor of anthropology and chemistry at Western Illinois University. Her research focuses on relationships between humans and their environments, including cultural and biological adaptations surrounding ancient diet, health, and knowledge systems; genetic disorders such as Neurofibromatosis Type 1 (NF1); effects of environmental factors on the preservation of bones, plant remains, and the molecules within them; and the comparative utility of field and laboratory techniques to produce informative archaeological, nutritional, and forensic data.<\/p>\n<p>Dr. Alveshere earned her B.A. in anthropology at the University of Washington with an emphasis in archaeology and an undergraduate research focus on the analysis of skeletal remains and geoarchaeological deposits. At the University of Minnesota, she completed her Ph.D. in anthropology, with a minor in human genetics. Her graduate thesis investigated factors that influence the preservation and detection of DNA in ancient and forensic specimens.<\/p>\n<p>Dr. Alveshere also worked for several years as a forensic scientist in the DNA\/Biology section of the Minnesota Bureau of Criminal Apprehension Forensic Science Laboratory. She led the WIU Archaeological Field School, on alternate summers since 2017, and conducted archaeological excavations in Israel, South Africa, and throughout the midwestern United States.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-589\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Stephanie-Etting-Author-Profile.jpg\" alt=\"\" width=\"201\" height=\"302\" \/><\/p>\n<h2>Stephanie Etting, Ph.D.<\/h2>\n<h3>(Writer Chapter 6)<\/h3>\n<p>Sacramento City College and Sonoma State University, ettings@scc.losrios.edu<\/p>\n<p>Dr. Etting became hooked on biological anthropology as a freshman at UC Davis when she took the \u201cIntroduction to Biological Anthropology\u201d course. She obtained her Ph.D. in anthropology in 2011 from UC Davis, where she studied anti-predator behavior toward snakes in rhesus macaques, squirrel monkeys, and black-and-white ruffed lemurs. While in graduate school, Dr. Etting discovered her love of teaching and, since finishing her dissertation, has taught at UC Berkeley; Sonoma State University; UC Davis; California State University, Sacramento; and Sacramento City College.In addition to her interests in primate behavior, Dr. Etting is also very interested in primate evolution and functional anatomy.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-590\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Karin-Enstam-Jaffe-Author-Profile.png\" alt=\"\" width=\"256\" height=\"200\" \/><\/p>\n<h2>Karin Enstam Jaffe, Ph.D.<\/h2>\n<h3>(Writer Chapter 7)<\/h3>\n<p>Sonoma State University, karin.jaffe@sonoma.edu<\/p>\n<p>Dr. Karin Enstam Jaffe has loved primates since she was five years old. As an undergraduate at U.C. San Diego, she participated in projects studying orangutans, langurs, and Mona monkeys. She earned her Ph.D. in Anthropology from U.C. Davis studying vervet and patas monkey antipredator behavior in Kenya. She has been a faculty member in the Anthropology Department at Sonoma State University since August 2002. A dedicated teacher-scholar, Dr. Jaffe has won several teaching, scholarship, and mentoring awards, including SSU\u2019s Excellence in Teaching Award, Educational Experience Enhancement Award, and the President\u2019s Excellence in Scholarship Award. In addition to teaching, she has led student research projects on behavioral enrichment involving ring-tailed lemurs, chimpanzees, and sun bears, as well as a study of the social network of hamadryas baboons at Oakland Zoo.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-592\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Sarah-S.-King-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"280\" \/><\/p>\n<h2>Sarah S. King, Ph.D.<\/h2>\n<h3>(Writer Chapter 8)<\/h3>\n<p>Cerro Coso Community College, sarah.king1@cerrocoso.edu<\/p>\n<p>Dr. Sarah S. King is an anthropology\/sociology professor at Cerro Coso Community College in California. She completed her Ph.D. work at the Division of Archaeological, Geographical and Environmental Sciences at the University of Bradford in West Yorkshire, England. Her thesis was entitled \u201cWhat Makes War?: Assessing Iron Age Warfare through Mortuary Behavior and Osteological Patterns of Violence.\u201d She also holds anthropology degrees from the University of California, Santa Cruz (B.A. hons., 2004), and the University of New Mexico, Albuquerque (M.A., 2006).<\/p>\n<p>&nbsp;<\/p>\n<h3><img class=\"alignnone wp-image-593\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Kara-Jones-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"282\" \/><\/h3>\n<h2>Kara Jones, M.A.<\/h2>\n<h3>(Writer Chapter 8)<\/h3>\n<p>PhD student at University of Nevada, Las Vegas, jonesk44@unlv.nevada.edu<\/p>\n<p>Kara Jones received their B.A. in anthropology at California State University, Bakersfield (2018) and their M.A. from University of Nevada, Las Vegas (2023, summer). Their master\u2019s thesis is titled \u201cRockin\u2019 at the Lake: Toolstone Use and Procurement along Holocene Lake Ivanpah, CA.\u201d Mx Jones is a Mojave Desert archaeologist specializing in stone tool use and manufacture, focusing further on Holocene lakeshore adaptations.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-594\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Jonathan-M.-G.-Perry-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"254\" \/><\/p>\n<h2>Jonathan M. G. Perry, Ph.D.<\/h2>\n<h3>(Writer Chapter 9)<\/h3>\n<p>Western University of Health Sciences, Oregon, jperry@westernu.edu<\/p>\n<p>Jonathan Perry was trained as a paleontologist and primatologist at the University of Alberta, Duke University, and Stony Brook University. His research focuses on the relationship between food, feeding, and craniodental anatomy in primates both living and extinct. This work includes primate feeding behavior, comparative anatomy, biomechanics, and field paleontology. He has taught courses on primate evolution at the undergraduate and graduate level.<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-595\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Stephanie-L.-Canington-Author-Profile.png\" alt=\"\" width=\"201\" height=\"183\" \/><\/p>\n<h2>Stephanie L. Canington, Ph.D.<\/h2>\n<h3>(Writer Chapter 9)<\/h3>\n<p>University of Pennsylvania, scaning@upenn.edu<\/p>\n<p>Stephanie Canington is a postdoctoral researcher at the University of Pennsylvania. Her current research is on the links between food properties, feeding behavior, and jaw morphology in lemurs that live in varying forms of captivity.<\/p>\n<p>&nbsp;<\/p>\n<h3><img class=\"alignnone wp-image-596\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Kerryn-Warren-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"200\" \/><\/h3>\n<h2>Kerryn Warren, Ph.D.<\/h2>\n<h3>(Writer Chapter 10)<\/h3>\n<p>Grad Coach International, kerryn.warren@gmail.com<\/p>\n<p>Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate Homo naledi remains from the Rising Star Cave System in the Cradle of Humankind.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-597\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/K.-Lindsay-Hunter-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"196\" \/><\/p>\n<h2>K. Lindsay Hunter, M.A., Ph.D. candidate<\/h2>\n<h3>(Writer Chapter 10)<\/h3>\n<p>CARTA, k.lindsay.hunter@gmail.com<\/p>\n<p>Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-598\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Navashni-Naidoo-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"230\" \/><\/p>\n<h2>Navashni Naidoo, M.Sc.<\/h2>\n<h3>(Writer Chapter 10)<\/h3>\n<p>University of Cape Town, nnaidoo2@illinois.edu<\/p>\n<p>Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-599\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Silindokuhle-Mavuso-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"200\" \/><\/p>\n<h2>Silindokuhle Mavuso, M.Sc.<\/h2>\n<h3>(Writer Chapter 10)<\/h3>\n<p>University of Witwatersrand, S.muvaso@ru.ac.za<\/p>\n<p>Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-600\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Bonnie-Yoshida-Levine-Author-Profile.jpg\" alt=\"\" width=\"201\" height=\"302\" \/><\/p>\n<h2><strong>Bonnie Yoshida-Levine, Ph.D. <\/strong><\/h2>\n<h3><strong>(Writer Chapter 11)\u00a0<\/strong><\/h3>\n<p>Grossmont College, bonnie.yoshida@gcccd.edu<\/p>\n<p>Bonnie Yoshida-Levine is an instructor of anthropology at Grossmont College, where she teaches biological anthropology and archaeology. She received her bachelor\u2019s degree in history from the University of California, Los Angeles, and her M.A. and Ph.D. degrees in anthropology from the University of California, Santa Barbara. Her dissertation research focused on the bioarchaeology of early civilizations in north coastal Peru. Bonnie has also collaborated on archaeological field projects in Bolivia and coastal California.<\/p>\n<p>&nbsp;<\/p>\n<h3><img class=\"alignnone wp-image-601\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Amanda-Wolcott-Paskey-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"280\" \/><\/h3>\n<h2>Amanda Wolcott Paskey, M.A.<\/h2>\n<h3>(Writer Chapter 12)<\/h3>\n<p>Cosumnes River College, paskeya@crc.losrios.edu<\/p>\n<p>Amanda Wolcott Paskey is an anthropology professor at Cosumnes River College in Sacramento, California. She earned her B.A. and M.A. in anthropology from the University of California, Davis. Her speciality in anthropology is archaeology; however, she was trained in a holistic program and most of her teaching load is in biological anthropology. She is currently working on analyzing a post\u2013gold rush era archaeological site, in Sacramento, with colleagues and students. This project has given her many opportunities to engage in sharing archaeology with a public audience, including local school children and Sacramentans interested in local history.<\/p>\n<h3><img class=\"alignnone wp-image-602\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/AnnMarie-Beasley-Cisneros-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"266\" \/><\/h3>\n<h2>AnnMarie Beasley Cisneros, M.A.<\/h2>\n<h3>(Writer Chapter 12)<\/h3>\n<p>American River College, beaslea@arc.losrios.edu<\/p>\n<p>AnnMarie Beasley Cisneros is an anthropology professor at American River College in Sacramento, California. Trained as a four-field anthropologist, she earned her B.A. and M.A. in anthropology from California State University, Sacramento. She regularly teaches biological anthropology, among other courses, and is currently engaged in applied anthropology work in community development with historically underserved communities. She most recently has particularly enjoyed facilitating her students\u2019 involvement in projects serving Sacramento\u2019s Latino and immigrant Mexican populations.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-603\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Keith-Chan-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"266\" \/><\/p>\n<h2>Keith Chan, Ph.D.<\/h2>\n<h3>(Writer Chapter 13)<\/h3>\n<p>Grossmont-Cuyamaca Community College District and MiraCosta College, drkeithcchan@gmail.com, Dr. Keith Chan is an instructor of anthropology at Grossmont-Cuyamaca Community College District and MiraCosta College in San Diego County. He reached this step of his anthropological path after many memorable experiences across the country and the hemisphere. He earned a bachelor\u2019s degree in anthropology from the University of California, Berkeley, in 2001. As a graduate student at the University of Missouri, he traveled to Per\u00fa with teams of students to study skeletons in the archaeological record to understand the lives of ancient Andeans. He completed his dissertation and earned a Ph.D. in 2011. Inspired by many educators in his journey, Dr. Chan turned his career toward teaching anthropology and helping students understand and appreciate humanity.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-604\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Michael-B.-C.-Rivera-Author-Profile.png\" alt=\"\" width=\"200\" height=\"208\" \/><\/p>\n<h2>Michael B. C. Rivera, Ph.D.<\/h2>\n<h3>(Writer Chapter 14)<\/h3>\n<p>University of Hong Kong, mrivera@hku.hk<\/p>\n<p>Michael B. C. Rivera is a biological anthropologist and human bioarchaeologist who studies human evolution and history and works to develop these disciplines in Hong Kong, East\/Southeast Asia, and the \u201cGlobal South.\u201d His doctoral thesis focused on the transition into agriculture in coastal environments and adaptations of ancient people along the beach. He is the only biological anthropologist working at the University of Hong Kong and the lead archaeologist managing the excavation of a WWII military aircraft that crashed in Hong Kong in 1945. Michael is also an advocate for greater inclusion, diversity, equality, and access to learning in academia. Much of his work also includes science communication and public engagement activities online, in schools, and in collaboration with museums.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-605\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Leslie-E.-Fitzpatrick-Author-Profile.jpg\" alt=\"\" width=\"266\" height=\"200\" \/><\/p>\n<h2>Leslie E. Fitzpatrick, Ph.D., RPA<\/h2>\n<h3>(Writer Chapter 15)<\/h3>\n<p>Independent Archaeological Consultants<\/p>\n<p>Lfitzpatrick@iac-llc.net<\/p>\n<p>Leslie Fitzpatrick is an historical archaeologist with Independent Archaeological Consultants based in Dover, New Hampshire. She earned a PhD in Anthropology from the University of Wyoming (2017), an MA in Anthropology from Georgia State (2012), and a BS in Mechanical Engineering from Georgia Tech (2000). Her primary research focus is the stable-isotope analysis of human remains as a means of interpreting past mobility and diet profiles for both modern and archaeological populations. In addition to her work as a historical archaeologist in New England, she has worked as a bioarchaeologist at field sites in Germany, Spain, Croatia, Mexico, Peru, and throughout the United States.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-606\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Ashley-Kendell-Author-Profile.jpg\" alt=\"\" width=\"201\" height=\"268\" \/><\/p>\n<h2>Ashley Kendell, Ph.D.<\/h2>\n<h3>(Writer Chapter 16)<\/h3>\n<p>California State University, Chico, akendell@csuchico.edu<\/p>\n<p>Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-607\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Alex-Perrone-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"266\" \/><\/p>\n<h2>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/h2>\n<h3>(Writer Chapter 16)<\/h3>\n<p>Butte Community College, perroneal@butte.edu<\/p>\n<p>Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-608\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Colleen-Milligan-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"266\" \/><\/p>\n<h2>Colleen Milligan, Ph.D.<\/h2>\n<h3>(Writer Chapter 16)<\/h3>\n<p>California State University, Chico, cfmilligan@csuchico.edu<\/p>\n<p>Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"alignnone wp-image-609\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Joylin-Namie-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"355\" \/><\/p>\n<h2>Joylin Namie, Ph.D.<\/h2>\n<h3>(Writer Chapter 17)<\/h3>\n<p>Truckee Meadows Community College, jnamie@tmcc.edu<\/p>\n<p>Joylin Namie is Professor of Anthropology at Truckee Meadows Community College, where she teaches courses in biological and cultural anthropology. Her current research interest is in culturally and environmentally sustainable tourism in desert environments, particularly in the country of Jordan and the U.S. state of Nevada. She was awarded a fellowship to Jordan from the Council of American Overseas Research Centers (CAORC) in 2020 to explore this topic, including visiting Petra and other important tourism destinations in Jordan. Dr. Namie\u2019s favorite things in life are teaching, competing in sports, and traveling.<\/p>\n<p>&nbsp;<\/p>\n<h3><\/h3>\n<h3><img class=\"alignnone wp-image-610\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Jonathan-Marks-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"216\" \/><\/h3>\n<h2>Jonathan Marks, Ph.D.<\/h2>\n<h3>(Writer Chapter 4)<\/h3>\n<p>University of North Carolina at Charlotte, jmarks@uncc.edu<\/p>\n<p>Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled What It Means to Be 98% Chimpanzee and Why I Am Not a Scientist, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p>&nbsp;<\/p>\n<h2><img class=\"alignnone wp-image-582\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2025\/08\/Adam-P-Johnson-Author-Profile.jpg\" alt=\"\" width=\"200\" height=\"200\" \/><\/h2>\n<h2>Adam P. Johnson, M.A.<\/h2>\n<h3>(Writer Chapter 4)<\/h3>\n<p>University of North Carolina at Charlotte\/University of Texas at San Antonio, ajohn344@uncc.edu<\/p>\n<p>Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<p>&nbsp;<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_958\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_958\"><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_1675_956\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_956\"><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 centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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 reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_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_1675_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 16.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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_1675_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 caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 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_1675_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1675_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_1675_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_1675_1178\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1178\"><div tabindex=\"-1\"><p>The role of a species in its environment; how it meets its needs for food, shelter, etc.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1180\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1180\"><div tabindex=\"-1\"><p>Competition that does not involve physical interaction between individuals, such as eating food before another individual arrives at the food site.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1182\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1182\"><div tabindex=\"-1\"><p>Organisms whose diet consists primarily of animal tissue.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1184\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1184\"><div tabindex=\"-1\"><p>The process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1012\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1012\"><div tabindex=\"-1\"><p>DNA molecule that is wrapped around protein complexes, including histones.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1010\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1010\"><div tabindex=\"-1\"><p>Proteins that DNA wraps around to assist with DNA organization within the nucleus.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1186\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1186\"><div tabindex=\"-1\"><p>An organism that lives in or on another organism.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1188\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1188\"><div tabindex=\"-1\"><p>A form of tourism that focuses on nature-based attractions to provide learning opportunities and that uses economically and ecologically sustainable practices.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1190\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1190\"><div tabindex=\"-1\"><p>A term used to describe females who are ready for sexual reproduction (i.e., not pregnant or nursing).<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1192\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1192\"><div tabindex=\"-1\"><p>The area that a group or individual uses over a given period of time (often over a year).<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1194\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1194\"><div tabindex=\"-1\"><p>The ability to avoid detection by other organisms, such as predators.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1196\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1196\"><div tabindex=\"-1\"><p>Active at night.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1198\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1198\"><div tabindex=\"-1\"><p>Watchful behavior used to detect potential danger, usually in the form of predators or potential competitors.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1200\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1200\"><div tabindex=\"-1\"><p>Vocalizations emitted by social animals in response to danger.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1202\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1202\"><div tabindex=\"-1\"><p>A way of describing which male(s) and female(s) mate.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1204\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1204\"><div tabindex=\"-1\"><p>Refers to the way in which animals move about their environment.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1206\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1206\"><div tabindex=\"-1\"><p>To leave one\u2019s group or area. This may or may not involve joining another group.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1208\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1208\"><div tabindex=\"-1\"><p>Nonaggressive social interactions and associations between individuals.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1210\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1210\"><div tabindex=\"-1\"><p>A temporary alliance between individuals.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1212\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1212\"><div tabindex=\"-1\"><p>The ranked organization of individuals established by the outcome of aggressive-submissive interactions.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1410\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1410\"><div tabindex=\"-1\"><p>A form of quadrupedal movement used by Gorilla and Pan when on the ground, wherein the front limbs are supported on the knuckles of the hands.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1412\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1412\"><div tabindex=\"-1\"><p>How the first living organism came into being.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1414\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1414\"><div tabindex=\"-1\"><p>The ancient organism from which all living things on Earth are descended.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1416\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1416\"><div tabindex=\"-1\"><p>Molecules that carry collections of genes.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1418\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1418\"><div tabindex=\"-1\"><p>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<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1420\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1420\"><div tabindex=\"-1\"><p>A sequence of DNA that provides coding information for the construction of proteins.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1422\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1422\"><div tabindex=\"-1\"><p>The entire collection of genetic material in a breeding community that can be passed on from one generation to the next.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1424\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1424\"><div tabindex=\"-1\"><p>A genotype comprising two different alleles.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1426\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1426\"><div tabindex=\"-1\"><p>The ratio, or percentage, of one allele compared to the other alleles for that gene within the study population.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1428\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1428\"><div tabindex=\"-1\"><p>A change in the nucleotide sequence of the genetic code. This is one of the forces of evolution.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1430\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1430\"><div tabindex=\"-1\"><p>Mutations that produce some sort of an advantage to the individual.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1432\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1432\"><div tabindex=\"-1\"><p>Enzymes that patrol and repair DNA in living cells.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1434\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1434\"><div tabindex=\"-1\"><p>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<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1436\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1436\"><div tabindex=\"-1\"><p>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<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1438\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1438\"><div tabindex=\"-1\"><p>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<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1675_1440\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1675_1440\"><div tabindex=\"-1\"><p>A point mutation that causes a change in the resulting protein.<\/p>\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":12,"template":"","meta":{"pb_show_title":"on","pb_short_title":"","pb_subtitle":"","pb_authors":["stephanie-etting"],"pb_section_license":""},"chapter-type":[],"contributor":[89],"license":[],"class_list":["post-1675","chapter","type-chapter","status-publish","hentry","contributor-stephanie-etting"],"part":20,"_links":{"self":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1675","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\/1675\/revisions"}],"predecessor-version":[{"id":1776,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1675\/revisions\/1776"}],"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\/1675\/metadata\/"}],"wp:attachment":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/media?parent=1675"}],"wp:term":[{"taxonomy":"chapter-type","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapter-type?post=1675"},{"taxonomy":"contributor","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/contributor?post=1675"},{"taxonomy":"license","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/license?post=1675"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}