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Essentials of Physical Anthropology, Anthropology, Eighth Eighth Edition Robert Jurmain, Lynn Kilgore, and Wenda Trevanthan Anthropology Editor: Erin Mitchell Developmental Editor: Lin Gaylord Assistant Editor: Rachael Krapf Editorial Assistant: Pamela Simon Media Editor: Melanie Cregger
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Focus Questions What do physical anthropologists do? Why is physical anthropolog y a scientific discipline, and what is its importance to the general public?
Dr. Russell Mittermeier, president
of Conservation International, holding an indri (the largest of the living lemurs) at a wildlife reser ve in Madagascar.
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Introduction to Physical Anthropology
1
One day, perhaps during the rainy season some 3.7 million years ago, two or three animals walked across a grassland savanna (see next page for definitions) in what is now northern Tanzania, in East Africa. These individuals were early hominins, members of the evolutionary lineage that includes ourselves, modern Homo sapiens. Fortunately for us, a
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record of their passage on that long-forgotten day remains in the form of footprints, preserved in hardened volcanic deposits. As chance would have it, shortly after heels and toes were pressed into damp soil, a nearby volcano erupted. The ensuing ashfal l blanketed everything on the ground, including the hominin footprints. In time, the ash layer hardened into a deposit that preserved the tracks for nearly 4 million years (Fig. 1-1). These now famous prints indicate that two individuals, one smaller than the other, may have walked side by side, leaving parallel sets of tracks. But because the larger individual’s prints are obscured, possibly by those of a third, it’s unclear how many actually made that journey so long ago. But what is clear is that the prints were made by an animal that habitually walked bipedally (on two feet), and that fact tells us that those ancient travelers were hominins. In addition to the footprints, scientists working at this site (called Laetoli) and at other locations have discovered many fossilized parts of skeletons of an an imal we call Australopithecus afarensis. After analyzing these remains, we know that these hominins were anatomically similar to ourselves, although their brains were only about one-third the size of ours. And even though they may have used stones and sticks as simple tools, there’s no evidence to suggest that they actually made stone tools. In fact, they were very much at the mercy of nature’s whims. They certainly couldn’t outrun most predators, and since their canine teeth were fairly small, they were pretty much defenseless. We’ve asked hundreds of questions about the Laetoli hominins, but we’ll never be able to answer them all. They walked down a path into what became their future, and their immediate journey has long since ended. So it remains for us to learn as much as we can about them and their species; and as we continue to do so, their greater journey continues. 3
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CHAPTER 1 Introduction to Physical Anthropology
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savanna (also spelled savannah) A large flat grassland with scattered trees and shrubs. Savannas are found in many regions of the world with dry and warmto-hot climates. hominins Colloquial term for members of the evolutionary group that includes modern humans and extinct bipedal relatives. bipedally On two feet; habitually walking on two legs. species A group of organisms that can interbreed to produce fertile offspring. Members of one species are reproductively isolated from members of all other species (that is, they cannot mate with them to produce fertile offspring). primates Members of the order of mammals Primates (pronounced “prymay´-tees”), which inclu des lemurs, lorises, tarsiers, monkeys, apes, and humans.
On July 20, 1969, a television audience numbering in the hundreds of millions watched as two human beings stepped out of a spacecraft onto the surface of the moon. To anyone born after that date, this event may be more or less taken for granted. But the significance of t hat first moonwalk can’t be overstated, because it represents humankind’s presumed mastery over the natural forces that govern our presence on earth. For the first time ever, people had actually walked upon the surface of a celestial body that, as far as we know, has never given birth to biological life. As the astronauts gathered geological specimens and f rolicked in near weightlessness, they left traces of their fleeting presence in the form of footprints in the lunar dust (Fig. 1-2). On the surface of the moon, where no rain falls and no wind blows, the footprints remain undisturbed to this day. They survive as mute testimony to a brief visit by a medium-sized, big-brained creature who presumed to challenge the very forces that created it. You may be wondering why anyone would care about early hominin footprints and how they can possibly be relevant to your life. You may also wonder why a physical anthropology textbook would begin by discussing two such seemingly unrelated events as hominins walki ng across a savanna and a moonwalk. But the fact is, these two events are closely related. Physical, or biological, anthropology is a scientific discipline concerned with the biological and behavioral characteristics of human beings; our closest relatives, the nonhuman primates (apes, monkeys, tarsiers, lemurs and lorises); and their ancestors. This kind of research helps us explain what it means to be human. This is an ambitious goal, and it probably isn’t fully attainable, but it’s certainly worth pursuing. Af ter all, we’re the only species to ponder our own existence and question how we fit into the spectrum of life on earth. Most people view humanity as separate from the rest of the anima l kingdom. But at the same s e n time, some are curious about the similarities we share with other species. Maybe, J o r e t as a child, you looked at your dog and tried to figure out how her front legs might P e correspond to your arms. Perhaps, during a visit to the zoo, you noticed the similarities between a chimpanzee’s hands or facial expressions and your own. Maybe you wondered if they also shared your thoughts and feelings. If you’ve ever had thoughts and questions like these, then you’ve indeed been curious about humankind’s place in nature. We humans, who can barely comprehend a century, can’t begin to grasp the enormity of nearly 4 million years. But we still want to know more about those creatures who walked across the savanna that day. We want to know how an insignif icant but clever bipedal primate such as Australopithecus afarensis, or perhaps a close relative, gave rise to a species that would eventually walk on the surface of the moon, some 230,000 miles from earth. How did Homo sapiens , a result of the same evolu-
FIGURE 1-1
Early hominin footprint s at Laetoli, Tanzania. The tracks to the left were made by one individual, while those to t he right appear to have be en formed by two individuals, the second stepping in the tracks of the first.
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tionary forces that produced all other life on this planet, gain the power to control the flow of rivers and even alter the climate on a global scale? As tropical animals, how were we able to leave the tropics and eventually occupy most of the earth’s land surfaces? How did we adjust to different environmental conditions as we dispersed? How could our species, which numbered fewer than 1 billion until the midnineteenth century, come to number almost 7 billion worldwide today and, as we now do, add another billion people every 11 years? These are some of the many ques FIGURE 1-2 tions that physical anthropologists A Human footprints left on the lunar S A N surface during the Apollo mission. attempt to answer through the study of human evolution, variation, and adaptation. These issues, and many others, are the topics covered directly or indirectly i n this textbook, because physical anthropology is, in large part, human biology seen from an evolutionary perspective. As biological organisms, human s are subjected to the same evolutionary forces as all other species a re. On hearing the term evolution, most people think of the appearance of new species. Certainly, the development of new species is one important consequence of evolution; but it isn’t the only one, because evolution is an ongoing biological process w ith more than one outcome. Simply stated, evolution is a change in the genetic makeup of a population from one evolution A change in the genetic generation to the next, and it can be defined and st udied at two levels. Over structure of a population. The term is time, some genetic changes in populations do result in the appearance of a new also sometimes used to refer to the species or speciation , especially when those populations are isolated from one appearance of a new species. another. Change at this level is cal led macroevolution . At the other level, there adaptation An anatomical, physiologare genetic alterations within populations; and while this type of change may not ical, or behavioral response of organisms lead to speciation, it often causes populations of a species to differ f rom one or populations to the environment. another regarding the frequency of certai n traits. Evolution at this level is Adaptations result from evolutionary referred to as microevolution. Evolution as it occurs at both these le vels will be change (specifically, as a result of natur al addressed in this book. selection). But biological anthropologists don’t just study physiological and biological genetic Pertaining to genetics, the systems. When these topics are considered within the broader context of human study of gene structure and action and evolution, another factor must be considered, and that factor is culture. Culture is the patterns of transmission of traits an extremely important concept, not only as it relates to modern human beings from parent to offspring. Genetic mechabut also because of its critical role in human evolution. Quite simply, and in a nisms are the foundation for evolution very broad sense, culture can be said to be the strategy by which people adapt to ary change. the natural environment. In fact, culture has so altered and so dominated our culture Behavioral aspects of human world that it’s become the environment in which we live. Culture includes technoladaptation, including technology, tradiogies ranging from stone tools to computers; subsistence patterns, from hunting tions, language, religion, marriage patand gathering to global agribusiness; housing types, from thatched huts to skyterns, and social roles. Culture is a set of scrapers; and clothing, from animal skins to high-tech synthetic fibers (Fig. 1-3). learned behaviors transmitted from one generation to the next by nonbiological Technology, religion, values, social organization, language, kinship, marriage rules, (that is, nongenetic) means. dietary preferences, gender roles, inheritance of property, and so on, are all aspects of culture. And each culture shapes people’s perceptions of the external envi ron worldview General cultural orientament, or their worldview , in particular ways that distinguish one culture from tion or perspective shared by members of all others. a society.
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CHAPTER 1
Introduction to Physical Anthropology
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FIGURE 1-3
Traditional and recent technolog y. (a) An early stone tool from East Africa. This artifact represents one of the oldest types of stone tools found anywhere. (b) The Hubble Space telescope, a late twentieth-century tool, orbits the earth every 96 minutes at an altitude of 360 miles. Because it is above the earth’s atmosphere, it provides distortion-free images of objects in deep space. (c) Cuneiform, the earliest form of writing, involved pressing symbols into clay tablets. It originated in southern Iraq some 5,000 years ago. (d) Text messaging, the most recent innovation in satellite communication, has generated a new language of sor ts. Currently more than 500 billion text messages are sent ever y day worldwide. (e) A Samburu woman in East Africa building a traditional but complicated dwe lling of stems, small branches, mud, and cow dung. (f ) These Hong Kong skyscrapers are typical of cities in industrialized countries today.
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One basic point to remember is that culture isn’t genetically pa ssed from one generation to the next. Culture is learned , and the process of learning one’s culture begins, quite literally, at birth. All humans are products of the culture they’re raised in, and since most of human behavior is learned, it follows that most behaviors, perceptions, and reactions are shaped by culture. At the same time, however, it’s important to emphasize that even though cultu re isn’t genetically determined, the human predisposition to assimilate culture and function within it is profoundly influenced by biological factors. Most nonhuman animals, including birds and especially primates, rely to varying degrees on learned behavior. This is especially true of the great apes (gorillas, chimpanzees, bonobos, and orangutans), which, as you will learn later, exhibit numerous aspects of culture. We can’t overemphasize that the predisposition for culture is perhaps the most critical component of human evolutionary history, and it was inherited from early hominin or prehominin ancestors. In fact, the common ancestor we share with chimpanzees may have had this predisposition. But during the course of human evolution, the role of culture beca me increasingly important. Over time, culture influenced many aspects of our biological makeup; and in tu rn, aspects of biology influenced cultural practices. For this reason, humans are the result of long-term interactions between biology and culture, and we call these interactions biocultural evolution. Biocultural interactions have resulted in many anatomical, biological, and behavioral changes during the course of human evolution: the shape of the pelvis and hip, increased brain size, reorganization of neurological structures, decreased tooth size, and the development of language, to name a few. What’s more, biocultural interactions are as important today as they were in the past, especially with regard to human health and disease. A ir pollution and exposure to dangerous chemicals have increased the prevalence of respiratory disease and cancer. And while air travel has made it possible for people to travel thousands of miles in just a few hours, we aren’t the only species that can do this. Disease-causing organisms travel with their human hosts, ma king it possible for infectious diseases l ike flu to spread, literally within hours, across the globe. Human activities have changed the patterns of infectious di seases such as tuberculosis and malaria. A fter the domestication of nonhuman animals, close contact with chickens, pigs, and catt le greatly increased human exposure to some of the diseases these animals carry. Through this contact we’ve also changed the genetic makeup of disease-causing microorganisms. For example, the “swine f lu” virus that caused the 2009 pandemic actually contains genetic material derived from bacteria that infect three di fferent species: humans, birds, and pigs. Also, by consuming meat and milk from infected anima ls, humans can acquire tuberculosis from cattle. And because we’ve overused antibiotics, we’ve made some strains of tuberculosis resistant to treatment and even deadly. As you can see, the interactions between humans, domesticated animals, and disea se-carrying organisms are complex, and we’re a long way from understanding how these interactions impact the pattern and spread of human infectious disease. Whi le it’s clear that we humans have influenced the development and spread of infectious disease, we still don’t know the many ways that changes in in fectious disease patterns a re affecting human biology and behavior. Anthropological research in this one area alone is enormously important to biomedical studies, and there are many other critical topics t hat biological anthropologists explore. So how does biological anthropology differ from human biology? In many ways it doesn’t, because human biologists also study human physiology, genetics, and adaptation. But human biology, as a discipline, doesn’t include studies of nonhuman primates or human evolution. So when biological research includes these topics as well as the role of culture in shaping our species, it’s placed within the discipline of anthropology .
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behavior Anything organisms do that involves action in response to internal or exter nal stimuli. The response of an individual, group, or species to its environment. Such response s may or may not be deliberate, and they aren’t necessarily the results of conscious decisi on making (for example, the behavior of one-celled organisms and insect s). biocultural evolution The mutual, interactive evolution of human biology and culture; the concept that biology makes culture possible and that developing culture further influences the direction of biological evolut ion; a basic concept in understanding the unique components of human evolution. anthropology The field of inquiry that studies human culture and evolutionary aspects of human biology; includes cultural anthropology, archaeology, linguistics, and physical, or biological, anthropology.
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CHAPTER 1 Introduction to Physical Anthropology
What Is Anthropology? Many anthropology majors are forced to contemplate thi s question when their friends or parents ask, “What are you studying?” The answer is often followed by a blank stare or a comment relating to Indiana Jones or dinosaurs. So, what is anthropology, and how is it different from sever al related disciplines? In the United States, anthropology is divided into four main subfields: cultural, or social, anthropology; linguistic anthropology; archaeology; and physical, or biological, anthropology. Each of these is divided into several special ized areas of interest. This four-field approach concerns all aspects of humanity across space and time. Each of the subdisciplines emphasizes different facets of humanity, but together, the four fields offer a means of explaining variation in human adaptations. In addition, each of these subfields has practical applications, and many anthropologists pursue careers outside the university environment. This kind of anthropology is called applied anthropology , and it’s extremely important today.
Cultural Anthropology
applied anthropology The practical application of anthropological and archaeological theories and techniques. For example, many biological anthropologists work in the public health sector. ethnographies Detailed descriptive studies of human societies. In cultural anthropology, an ethnography is traditionally the stud y of a non-Western society.
Cultural, or social, anthropology is the study of the global patterns of belief and behavior found in modern and historical cultures. The origins of cultural anthropology can be traced to the nineteenth century, when travel and exploration increasingly brought Europeans into contact (and sometimes conflict) with various cu ltures in Africa, par ts of Asia, and the South Pacific islands . Also, i n the New World, there was considerable interest in Native Americans. This interest in “traditional” societies led early a nthropologists to study and record lifeways that are now mostly extinct. These studies produced many descriptive ethnographies that covered a range of topics, including religion, ritual, myth, use of symbols, diet, technology, gender roles, and child-rearing practices. Ethnographic accounts, in turn, formed the basis for comparative studies of numerous cultures. By examini ng the similarities and dif ferences among cultures, cultura l anthropologists have been able to formulate many hypotheses regarding fundamental aspects of human behavior. The focus of cultura l anthropology shifted over the course of the t wentieth century. Cultural anthropologists still work in remote areas, but increasingly they’ve turned their gaze inward, toward their own countries and the people around them. Many contemporary cultura l anthropologists are concerned with the welfare of refugees and study their resettlement and cu ltural integration (or lack thereof) in the United States, Canada, and many European countries. Increasingly, ethnographic techniques have been applied to the study of diverse subcultures and their interactions with one another in contemporary metropolitan areas (urban anthropology). Medical anthropology is an applied subfield of cultur al anthropology that explores the relationship between various cultur al attributes and health and disease. One area of interest is how dif ferent groups view disease processes and how these views affect treatment or the willingness to accept treatment. When a medical anthropologist focuses on the social dimensions of disease, physicians and physical anthropologists may also collaborate. In fact, many medical anthropologists have received much of their training in public health or physical anthropology.
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Archaeology Archaeology is the study of earlier cultures and l ifeways by anthropologists who specialize in the scientific recovery, analysis, and interpretation of the material remains of past societies. Although archaeology often concerns cultures that existed before the invention of writing (the period known as prehistory), historic archaeologists study the evidence of later, more complex societies that produced written records. Archaeologists are concerned with culture, but instead of studying living people, they obtain information from artifacts and structures left behind by earlier cultures. The remains of earlier societies, in the form of tools, structures, art, eating implements, fragments of writing, and so on, provide a great deal of information about many important aspects of a society, such as religion and social structure. Unlike in the past, sites aren’t excavated simply for the artifacts or “treasures” they may contain. Rather, they’re excavated to gain information about human behavior. For example, patterns of behavior are ref lected in the dispersal of human settlements across a landscape and in t he distribution of cultural remains within them. Archaeological research may focus on specific local ities or peoples and attempt to identify, for example, various aspects of social organization or factors that led to the collapse of a civi lization. Alternatively, inquiry may ref lect an interest in broader issues relating to human cultu re in general, such as the development of agriculture or the rise of cities. In the United States, the greatest expansion in archaeology since the 1960s has been in the area of cultural resource management . This is an applied approach that arose from environmental legislation requiring a rchaeological evaluations and sometimes excavation of sites that may be threatened by development. (Canada and many European countries have similar legislation.) Many contract archaeologists (so called because their serv ices are contracted out to developers or contractors) are a ffiliated with private consulting f irms, state or federal agencies, or educational institutions. In fact, a n estimated 40 percent of all archaeologists in the United States now fill such positions.
Linguistic Anthropology Linguistic anthropology is the study of human speech and language, including the origins of language in general as well as specif ic languages. By examining similarities between contemporary languages, linguists have been able to trace historical ties between specific l anguages and groups of languages, thus facilitating the identification of language famil ies and perhaps past relationships between human populations. Because the spontaneous acquisition and use of language is a uniquely human characteristic, it’s an important topic for linguistic anthropologists, who, along with specialists in other fields, study the process of language acquisition in infants. Since insights into the process may well have implications for the development of language skills in human evolution, as well as in growing children, it’s also an important subject to physical anthropologists.
Physical Anthropology As we’ve already said, physical anthropology is the study of human biology within the framework of evolution and with an emphasis on the interaction between
artifacts Objects or materials made or modified for use by moder n humans and their ancestors. The earliest artifacts tend to be tools made of stone or, occasionally, bone.
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CHAPTER 1 Introduction to Physical Anthropology
paleoanthropology The interdisciplinary approach to the study of earlier hominins—their chronology, physical structure, archaeological remains, habitats, and so on. anthropometry Measurement of human body parts. When osteologists measure skeletal elements, the term osteometry is often used.
biology and culture. This subdiscipline is also referred to as biological anthropology, and you’ll find the terms used interchangeably. Physical anthropology is the original term, and it reflects the initial interests anthropologists had in describing human physical variation. The American Association of Physical Anthropologists, its journal, as well as many college courses and numerous publications, retain this term. The designation biological anthropology reflects the shift in emphasis to more biologically oriented topics, such as genetics, evolutionary biology, nutrition, physiological adaptation, and growth a nd development. This shift occurred largely because of advances in the field of genetics and molecular biology since the late 1950s. Although we’ve continued to use the traditional term in t he title of this textbook, you’ll find that all of the major topics pertain to biological issues. The origins of physical anthropology can be found in two principal areas of interest among nineteenth-century European and American scholars. Many scientists (at the time called natural historians or naturalists) became increasingly curious about the origins of modern species. They were beginning to doubt the literal interpretation of the biblical account of creation at a time when scientific explanations emphasizing natural processes rather t han supernatural phenomena were becoming more popular. Eventually, these sparks of interest in biological change over time were fueled into flames by the publication of Charles Darwin’s On the Origin of Species in 1859. Today, paleoanthropology , or the study of human evolution, as evidenced in the fossil record, is a major subfield of physical anthropology (Fig. 1-4). Thousands of specimens of human ancestors (mostly fragmentary) are now kept in research collections. Taken together, these fossils span about 7 million years of human prehistory; and although most of these fossils are incomplete, they provide us with a significant wealth of knowledge that increases each year. It’s the ultimate goal of paleoanthropological research to identify the various early human and human-like species, establish a chronological sequence of relationships among them, and gain insights into their adaptation and behavior. Only then wil l we have a clear picture of how and when humankind came into being. Human variation was the other major area of interest for early biological anthropologists. They were especially concerned with observable physical differences, skin color being the most obvious. Enormous effort was aimed at describing and explaining the biological differences between various human populations. Although some attempts were misguided and even racist, they gave birth to many body measurements that are sometimes still used. Physical anthropologists also use many of the techniques of anthropometry to study skeletal remains from archaeological sites (Fig. 1-5). Moreover, anthropometric techniques have had considerable application in the design of everything from wheelchairs to off ice furniture. Undoubtedly, they’ve also been used to determine the absolute minimum amount of leg room a person must have in order to complete a 3-hour f light on a commercial airliner and remain s ane. Today, anthropologists are concerned with human variation because of its possible adaptive significance and because they want to identify the genetic and other evolutionary factors that have produced variation. In other words, some traits that t ypify certain populations evolved as biological adaptations, or adjustments, to local environmental conditions such as sun light, altitude, or infect ious disease. Other traits may simply be the results of geographical isolation or the descent of populations from small founding groups. Examini ng biological variation between populations of any species provides valuable information as to the mechanisms of genetic change in groups over time, and this is really what the evolutionary process is all about. Modern population studies also examine other important aspects of human variation, including how different groups respond physiologically to different
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a
FIGURE 1-4
(a) Paleoanthropologists excavating at the Drimolen site, South Africa. (b) Primate paleontologist Russ Ciochon and Le Trang Kha, a vertebrate pale ontologist, examine the fossil remains of Gigantopithecus from a 450,000-yearold site in Vietnam. Gigantopithecus is the name given to the largest apes that ever lived. In the background is a reconstruction of this enormous animal.
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FIGURE 1-5
Anthropology student using spreading calipers to measure cranial length.
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CHAPTER 1 Introduction to Physical Anthropology
FIGURE 1-6
This researcher is using a treadmill test to assess a subject’s heart rate, blood pressure, and oxygen consumption.
kinds of environmentally induced stress (Fig. 1-6). Such stresses may include high altitude, cold, or heat. Many biological anthropologists conduct nutritional studies, investigating the relationships between various dietary components, cultural practices, physiology, and certain aspects of health and disease (Fig. 1-7). Investigations of human fertility, growth, and development also are closely related to the topic of nutrition. r e n These fields of inquiry, which are i e t s n fundamental to studies of adapta e g e R tion in modern human popula h t i d u J tions, can provide insights into hominin evolution, too. It would be impossible to study evolutionary process es without some knowledge of how traits are inherited. For this reason and others, genetics is a crucial field for physical anthropologists. Modern physical anthropology wouldn’t exist as a n evolutionary science i f it weren’t for advances in t he understanding of genetic mechanisms. In this exciting time of rapid advances in genetic research, molecular anthro pologists use cutting-edge technologies to investigate evolutionary relationships between human populations as well as between humans and nonhuman primates. To do this, they examine similarities and differences in DNA sequences between
DNA (deoxyribonucleic acid) The double-stranded molecule that contains the genetic code. DNA is a main component of chromosomes.
FIGURE 1-7
Dr. Kathleen Galvin measures upper arm circumference in a young Maasai boy in Tanzania. Data derived from various body measurements, including height and weight, were used in a health and nutrition study of groups of Maasai cattle herders.
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FIGURE 1-8
(a) Cloning and sequencing method s are frequently used to identif y genes in humans and nonhuman primates. This graduate student identifies a genetically modified bacterial clone. (b) Molecular anthropologist Nelson Ting collecting red colobus fecal samples for a study of genetic variation in small groups of monkeys isolated from one another by agricultural clearing.
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individuals, populations, and species. What’s more, by extracting DNA from certain fossils, these researchers have contributed to our understanding of relationships between extinct and living species. As genetic technologies continue to be developed, molecular anthropologists will play a key role in explaining human evolution, adaptation, and our biological relationships with other species (Fig. 1-8). However, before genetic and molecular tech niques became widespread, osteology , the study of the skeleton, was the only way that anthropologists could study our immediate ancestors. In fact , a thorough knowledge of skeletal structure and function is stil l critical to the interpretation of fossil material today. For this reason, osteology has long been viewed as central to physical a nthropology. In fact, it’s so important that when many people think of biological anthropology, the first thing that comes to mind is bones! Bone biology and physiology are of major importance to many other aspects of physical anthropology besides human evolution. Many osteologists specialize in the measurement of skeletal elements, essential for identifying stature and growth patterns in archaeological populations. In the last 30 years or so, the study of human skeletal remains from archaeological sites has been cal led bioarchaeology . In turn, paleopathology , the study of disease and trauma in archaeologically derived skeletons, is a major component of bioarchaeology. Paleopathology is a prominent subfield that investigates the prevalence of trauma, certain infectious diseases (for instance, syphilis or tuberculosis), nutritional deficiencies, and numerous other conditions that m ay leave evidence in bone (Fig. 1-9). This research tells us a great deal about the lives of individuals and populations
osteology The study of skeletal material. Human osteology focuses on the interpretation of the skeletal remains from archaeological sites, skeletal anatomy, bone physiology, and growth and development. Some of the same techniques are used in paleoanthropology to study early hominins. bioarchaeology The study of skeletal remains from archaeological sites. paleopathology The study of disease and injury in human skeletal (or, occasionally, mummified) remains from archaeological sites.
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CHAPTER 1 Introduction to Physical Anthropology
a
FIGURE 1-9 Two examples of pathologi cal conditions in human skeletal remains from the Nubian site of Kulubnart i in Sudan. These remains are approximately 1,000 years old. (a) A partially healed fracture of a child’s left femur (thigh bone). The estimated age at death is 6 years, and the cause of death was probably an infection resulting from this injur y. (b) Very severe congenital scoliosis in an adult male from Nubia. The curves are due to developmental defects in individual vertebrae. (This is not the most common form of scoliosis.)
b
forensic anthropology An applied anthropological approach dealing with legal matters. Forensic anthropologists work with coroners and others in identifying and analyzing human remains.
FIGURE 1-10
Forensic anthropologists Vuzumusi Madasco (from Zimbabwe) and Patricia Bernardi (from Argentina) excavating the skeletal remains and clothing of a victim of a civil war massacr e in El Salvador. This burial is part of a mass grave, which was being excavated in order to try to identify vict ims and provide other information relative to the massacre.
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from the past. Paleopathology also yields information regarding the history of certain disease processes, and for this reason it’s of interest to scientists in biomedical fields. Forensic anthropology is directly related to osteology and paleopathology and has become of increasing interest to the public because of TV shows li ke Bones. Technically, this approach is the application of anthropological (usual ly osteological and sometimes archaeological) techniques to legal issues (Fig. 1-10). Forensic anthropologists help identify skeletal remains in mass disasters or other situations where a human body has been found. They’ve been involved in numerous cases having important legal, historical, and human consequences. They were also instrumental in identifying the skeletons of most of the Russian imperial family, executed in 1918. And many participated in the overwhelming task of trying to identify the remains of victims of the September 11, 2001, terrorist attacks in the United States. Anatomy is yet another important area of interest for physical anthropologists. In living organisms, bones and teeth are intimately linked to the soft tissues
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FIGURE 1-11
Dr. Linda Levitch teaching a human anatomy class at the University of North Carolina School of Medicine.
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that surround and act on them. Consequently, a thorough knowledge of soft tissue anatomy is essential to the understanding of biomechanical relationships involved in movement. Such relationships are important in accurately assessing the structure and f unction of limbs and other components of fossilized remains. For these reasons and others, many physical anthropologists specialize in anatomical studies. In fact, several physical anthropologists hold professorships in anatomy departments at universities and medical schools (Fig. 1-11). Primatology , the study of the living nonhuman primates, has become increasingly important since the late 1950s (Fig. 1-12). Behavioral studies, especially those conducted on groups in natura l environments, have implications for many scientific disciplines. Because nonhuman primates are our closest l iving relatives, identifying the underlying factors related to their social behavior, communication, infant care, reproductive behavior, and so on, helps us develop a better understanding of the natural forces that have shaped so many aspects of modern human behavior. It’s also very important to study nonhuman primates in their own right, regardless of what we may learn about ourselves. This is part icularly true today because the majority of primate species are threatened or seriously endangered. Only through research will scientists be able to recommend policies that can better ensure the survival of many nonhuman primates and thousands of other species as well. FIGURE 1-12
Primatologist Jill Pruetz follows a chimpanzee in Senegal, in West Africa.
primatology The study of the biology and behavior of nonhuman primates (prosimians, monkeys, and ap es).
k i n s e L e i l u J
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CHAPTER 1 Introduction to Physical Anthropology
Applied Anthropology Applied approaches in biological anthropology are numerous. And while applied anthropology is aimed at the practical application of anthropological theories and methods outside the academic setting, applied and academic anthropology aren’t mutually exclusive approaches. In fact, applied anthropology relies on the research and theories of academic anthropologists and at the same time has much to contribute to theory and techniques. Within biological anthropology, forensic anthropology is a good example of the applied approach. But the practical application of the techniques of physical anthropology isn’t new. During World War II, for example, physical anthropologists were extensively involved in designing gun turrets and airplane cockpits. Since then, many physical anthropologists have pursued careers in genetic and biomedical research, public health, evolutionary medicine, medical anthropology, and conservation of nonhuman primates, and many hold positions in museums and zoos. In fact, a background in physical anthropology is excellent preparation for almost any career in the medical and biological fields (Fig. 1-13).
a
b
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FIGURE 1-13
(a) Dr. Soo Young Chin, Lead Part ner of Practical Ethnogr aphics at Ascension Health, pointing to pilot locati ons for a study of a new health care plan. (b) Nanette Barkey, a medical anthropol ogist involved in a repatriation project in Angola, photographed this little girl being vaccinated at a refugee transit camp. Vaccinations were being administered to Angolan refugees returning home in 2004 from the Demo cratic Republ ic of Congo, where they had fled to escape warfare in their own country.
y e k r a B e t t e n a N
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From this brief overview, you can see that physical anthropology is the subdiscipline of anthropology that focuses on many aspects of human biology and evolution. Humans are a product of the same forces that produced all life on earth. As such, we’re just one contemporary component of a vast biological continuum at one point in time; and in this regard, we aren’t particularly unique. Stating that humans are part of a continuum doesn’t imply that we’re at the peak of development on that continuum. Depending on the criteria used, humans can be seen to exist at one end of the spectrum or the other, or somewhere in between, but we don’t necessarily occupy a position of inherent superiority over other species. However, human beings are truly unique in one dimension, and that is intellect. After all, humans are the only species, born of earth, to stir t he lunar dust. We’re the only species to develop language and complex culture as a means of buffering nature’s challenges; and by so doing, we have gained the power to shape the planet’s very destiny.
Physical Anthropology 17 and the Scientific Method
Physical Anthropology and the Scientific Method Science is an empirical approach to gaining information. It involves observing phenomena; developing hypotheses or possible explanations for them; and then
devising a research design or series of experiments to test those hypotheses. Because biological anthropologists are scientists, they adhere to t he principles of the scientific method by identifying a research problem and then gathering information to solve it. Once a question has been asked, the first step usually is to explore the existing literature (books and journals) to determine what other people have done to resolve the issue. Based on this preliminar y research and other observations, one or even several tentative explanations (hypotheses) are then proposed. The next step is to develop a research design or methodology aimed at testing the hypothesis. These methods involve collecting information or data that can then be studied and analyzed. Data can be a nalyzed in many ways, most of them involving various statistical tests. During the data collection and analysis phase, it’s important for scientists to use a rigorously controlled approach so they can precisely describe their techniques and results. This precision is critical because it enables others to repeat the experiments and allows scientists to make comparisons between their study and the work of others. For example, when scientists collect data on tooth size in hominin fossils, they must specify which teeth are being measured, how they’re measured, and the results of the measurements (expressed numerically, or quantitatively ). Then, by analyzing the data, the investigators try to draw conclusions about the meaning and significance of their measurements. This body of information then becomes the basis of future studies, perhaps by other researchers, who can compare their own results with those a lready obtained. Hypothesis testing is the very core of the scientific method, and although it may seem contradictory at first, it’s based on the potential to falsify the hypothesis. Falsification doesn’t mean that the entire hypothesis is untrue, but it does indicate that there may be exceptions to it. Thus, the hypothesis may need to be refined and subjected to further testing. Eventually, if a hypothesis stands up to repeated testing, it may become part of a theory , or perhaps a theory itself. There’s a popular misconception that a theory is mere conjecture, or a “hunch.” But in science, theories are proposed
continuum A set of relationships in which all components fall alo ng a single integrated spectrum (for example, color). All life reflects a single biological continuum. science A body of knowledge gained through observation and experimentation; from the Latin scientia, meaning “knowledge.” empirical Relying on experiment or observation; from the Latin empiricus, meaning “experience d.” hypotheses (sing., hypothesis) A provisional explanation of a phenomenon. Hypotheses require verification or falsification through testing. scientific method An approach to research whereby a problem is identified, a hypothesis (or provisional explanat ion) is stated, and that hypothesis is tested by collecting and analyzing data. data (sing., datum) Facts from which conclusions can be drawn; scientific information. quantitatively Pertaining to measurements of quantit y and including such propert ies as size, number, and capacity. When data are quantified, they’re expressed numerically and can be tested statistically. theory A broad statement of scientific relationships or underlying principles that has been substantially verified through the testing of hypotheses.
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CHAPTER 1 Introduction to Physical Anthropology
scientific testing The precise repetition of an experiment or expansion of observed data to provide verification; the procedure by which hypotheses and theories are verified, modified, or discarded.
explanations of relationships between natural phenomena. Theories usually concern a broader, more universal view than hypotheses, which have a narrower focus and deal with more specific relationships between phenomena. But like hypotheses, theories aren’t facts. They’re tested explanations of facts. For example, it’s a fact that when you drop an object, it falls to the ground. The explanation for this fact is the theory of gravity. Also, theories can be altered over time with further experimentation or observations as well as newly developed technologies. The theory of gravity has been tested many times and qual ified by experiments showing how the mass of objects affects how they’re attracted to one another. So far, the theory has held up. Scientific testing of hypotheses may ta ke several years (or longer) and may involve researchers who weren’t involved with the original work. What’s more, new methods may permit different kinds of testing that weren’t previously possible, and this is a strength of scientific research. For example, since the 1970s, primatologists have reported that male nonhuman primates (as well as males of many other species) sometimes kill infants. One hypothesis has been that these males were killing infants fathered by other males. Many scientists have objected to this hypothesis, and they’ve proposed several alternatives. For one thing, there was no way to know for certain that the males weren’t killing their own offspring; and if they were, this would argue against the hypothesis. However, in a fa irly recent study, scientists colle cted DNA samples from dead infants and the males who killed them and showed that most of the time, the males weren’t related to their victims. This result doesn’t prove that the original hypothesis is accurate, but it does strengthen it. This study is described in more detail i n Chapter 7, but we mention it here to emphasize that science is an ongoing process that builds on previous work and benefits from newly developed techniques (in this case, DNA testing) in ways that constantly expand our knowledge. Another current scientific debate focuses on how to interpret the remarkable small hominins found in Indonesia, popularly referred to in the media as “hobbits.” One hypothesis suggests that these smal l-bodied, small-brained hominins were members of a species other than Homo sapiens . A second hypothesis suggests that the remains a re those of modern humans with a pathological growth defect. As new methods a nd more intense analyses of the remains continue, these hypotheses are being tested, and we’ll discuss t he latest results in Chapter 11. There’s one more important fact about hypotheses and theories: Any proposition that’s stated as absolute and/or doesn’t allow the possibility of falsificati on is not a scientific hypothesis, and it should never be considered as such. We’ve emphasized that a crucial aspect of scientific statements is that there must be way to evaluate their validity. Statements such as “Heaven exists” may well be true (that is, they may describe some actual state), but there’s no rational, empirical means (based on experience or experiment) of testing them. Therefore, acceptance of such a view is based on faith rather than on scientific verification. The purpose of scientific research isn’t to establish absolute truths; rather, it’s to generate ever more accurate and consistent explanations of phenomena in our uni verse based on observation and testing. At its very heart, scientific methodolog y is an exercise in rational thought and critical thinking. The development of critical think ing skills is an important and lasting benefit of a college education. Such skills enable people to evaluate, compare, analyze, critique, and synthesize information so they won’t accept everything they hear at face value. Perhaps the most glaring need for critical thinking is in how we evaluate advertising claims. For example, people spend billions of dollars every year on
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“natural” dietary supplements based on marketing claims that may not have been tested. So when a salesperson tells you that, for example, echinacea helps prevent colds, you should ask if that statement has been scientifical ly tested, how it was tested, when, by whom, and where the results were published. Similarly, when politicians make claims in 30-second sound bytes, check those claims before you accept them as truth. Be skeptical, and if you do check the validity of advertising and political statements, you’ll find that frequently they’re either misleading or just plain wrong.
The Anthropological Perspective Perhaps the most important benefit you’ll receive from this textbook, and this course, is a wider appreciation of the human experience. To understand human beings and how our species came to be, we must broaden our viewpoint, through both time and space. All branches of anthropology fundamentally seek to do this in what we call the anthropological perspective . Physical anthropologists, for example, are interested in how humans differ from and are similar to other animal s, especially nonhuman primates. For example, we’ve defined hominins as bipedal primates, but what are the major anatomical components of bipedal locomotion, and how do they differ from, say, those in a quadrupedal ape? To answer these questions, we would need to study the anatomical structures involved in human locomotion (muscles, hips, legs, and feet) and compare them with the same structures i n various nonhuman primates. From a perspective that is broad in space and time, we can begin to grasp the diversity of the human experience within the context of biological and behavioral continuity with other species. In this way, we may better understand the l imits and potentials of humankind. Furthermore, by extending our knowledge to include cultures other than our own, we may hope to avoid the ethnocentric pitfalls inherent in a more limited view of humanity. This relativistic view of culture is perhaps more important now than ever before, because in our increasingly interdependent global community, it allows us to understand other people’s concerns and to view our own culture from a broader perspective. Likewise, by examining our species as part of a wide spectrum of life, we realize that we ca n’t judge other species using human criteria. Each species is unique, with needs a nd a behavioral repertoire not exactly like that of any other. By recognizing t hat we share many similarities (both biological and behavioral) with other animals, perhaps we may come to recognize that they have a place in nature just as surely as we ourselves do. In addition to broadening perspectives over space (that is, encompassing many cultures and ecological ci rcumstances as well as nonhuman species), an anthropological perspective al so extends our horizons through time. For example, in Chapter 17 we’ll discuss human nutrition. The vast majority of the foods people eat today (coming from domesticated plants and animals) were unavailable until around 10,000 years ago. Human physiological mechanisms for chewing and digesting foods nevertheless were already well established long before that date. These adaptive complexes go back millions of years. Besides the obviously different diets prior to the development of agriculture, earlier hominins might well have differed from humans today in average body size, metabolism, and activity patterns. How, then, does the basic evolutionary “equipment” (that is, physiology) inherited from our hominin forebears accommodate our modern diets? Clearly, the way to understand such processes is not just by looking at
ethnocentric Viewing other cultures from the inherently biased perspective of one’s own culture. Ethnocentrism often results in other cultures being seen as inferior to one’s own. relativistic Viewing entities as they relate to something else. Cultural relativism is the view that cultures have merits within their own historical and environmental contexts. metabolism The chemical processes within cells that break down nutrients and release energy for the body to use. When nutrients are broken down into their component part s, such as amino acids, energy is released and made available for use by the cell.
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CHAPTER 1 Introduction to Physical Anthropology
contemporary human responses, but by placing them in the perspective of evolutionary development through time. We hope that after reading the following pages, you’ll have an increased understanding not only of the similarities we share with other biological organisms but also of the processes that have shaped the traits that make us unique. We live in what may well be our planet’s most crucial period in the pa st 65 million years. We are members of the one species that, through the very agency of culture, has wrought such devastating changes in ecological systems that we must now alter our technologies or face potentially unspeakable consequences. In such a t ime, it’s vital that we attempt to gain the best possible understanding of what it means to be human. We believe that the study of physical anthropology is one endeavor that aids in this attempt, and that is indeed the goal of this textbook.
Why It Matters
T
oday, the trend in advanced education is toward greater and greater specialization, with the result that very few people or professions have the broad overview necessary to implement policy and make effective changes that could lead to improved standards of living, a safer geopolitical world, and better planetary health. This is acutely felt in medicine, where specialists focusing on one part of the body sometimes ignore other parts, often to the detriment of overall health (especially
mental and emotional) of the patient. Anthropology is one of the few disciplines that encourages a broad view of the human condition. An example is seen in AIDS prevention research. The wealth of knowledge that biologists and medical researchers have provided on the characteristics and behavior of HIV (the virus that causes AIDS) is useles s for preventing its transmission unless we also have an understanding of human behavior at both the individual and the sociocultural levels. Behavioral scientists, including anthropologists, are prepared to
examine the range of social, religious, economic, political, and historical contexts surrounding sexuality to devise AIDS prevention strategies that will vary from population to population and even from subculture to subculture. Whether or not you choose a career in anthropolo gy, the perspectives that you gain from studying this discipline will enable you to participate in research and policy decisions on future challenges to human and planetary health and well-being.
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Licensed to: iChapters User Critical Thinking Questions
Summary In this chapter, we’ve introduced you to the field of physical, or biological, anthropology, placing it within the overall context of anthropological st udies. As a major academic discipline within the social sciences, anthropology also includes cultural anthropology, archaeology, and linguistic anthropology as major subfields. Physical anthropology is the study of many aspects of human biolog y, including genetics, genetic variation, adaptations to environmental factors, nutrition, and anatomy. These topics are discussed within an evolutionary framework because all human characteristics are either directly or indirectly the results of biological evolution, which in turn is dr iven by genetic change. Hence, biological anthropologists also study our closest relatives, the nonhuman primates, primate evolution, and the genetic and fossil evidence for human evolution. Because biological anthropology is a scientific discipline, we also discussed the
role of the scientific method in re search. We presented the importance of objectivity, observation, data collection, and analysis; and we described the formation and testing of hypotheses t o explain natural phenomena. We also emphasized that this approach is an empirical one that doesn’t rely on supernatural explanations. Because evolution is the core of physical anthropology, in the next chapter we present a brief historical overview of changes in Western scientific thought that led to the discovery of the basic principles of biological evolut ion. As you’re probably aware, evolution is a highly controversial subject in the United States and increasingly in many Islamic countries. However, it’s not part icularly controversial in Europe. In the next chapter, we’ll address some of the reasons for this controversy and explain the evidence for evolution as the single thread uniting all the biological sciences.
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Critical Thinking Questions 1. Given that you’ve only just been introduced to the field of physical anthropology, why do you think subjects such as anatomy, genetics, nonhuman primate behavior, and human evolution are integrated into a discussion of what it means to be human? 2. Is it important to you, personally, to know about human evolution? Why or why not? 3. Do you see a connection between hominin footprints that are almost 4 million years old and human footprints left on the moon in 1969? If so, do you think this relationship is important? What does the fact that there are human footprints on the moon say about human adaptation? (You may wish to refer to both biological and cultural adaptation.)
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Licensed to: iChapters User o t o h p k c o t S i / k r i k s u B n a V e i m a J ©
Making a Difference: Forensic Anthropologists in the Contemporary World ue to wide media coverage, especially several popular television
Dshows, forensic anthropology has captured the i magination of many people. In addition to their well-known participation in assisting law enforcement official s investigating crime scenes, forensic anthropologists also work in a variety of other interesting situ ations. They are often called to join recovery teams at scenes of mass d isasters such a s the World Trade Center, plane crashes, or in areas devastated by an eart hquake or tsunami. Additionally, they’re involved in excavating mass g raves where victims of political atrocities have been secretly bur ied. These sites of such enormous human tragedy sadly a re found in many part s of the world, from Iraq to Bosnia, to Argentina, to Rwa nda, Forensic anthropologists also help search for and identif y soldiers missing in action from prior wars. In al l these difficult circu mstances, wherever possible, the goal is to determine the identity of missing people and to return their remains to family members. e c n a r F e n a i D ©
Scene of a Korean Airlines crash in
1996 in the U.S. terr itory of Guam, that killed 228 people. The U.S. government immediately sent numerous DMORT (Disaster Mortuary Operation Response) teams, each of which usually has at least one forensic anthropologist. All human remains were evaluated in
the field laborator y where Tom Holland (Director of the Central Identification Laborator y in Hawaii) is shown identifying fragmentar y skeletal elements, many of which were heavily burned (as were many of the bodies). Nevertheless, all the passengers and crew were accounted for. e c n a r F e n a i D ©
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e c i f f O n o s i a i L e m i r C e m i g e R e h t d n a , s r e e n i g n E f o s p r o C y m r A . S . U
Forensic anthropologists, including both physical anthropolo-
gists and archaeologists, re covered 114 Kurdish victims of genocide from this site in southern Iraq.
e c i f f O n o s i a i L e m i r C e m i g e R e h t d n a , s r e e n i g n E f o s p r o C y m r A . S . U
e d o o G . C k c i r r e D . t g S f f a t S y b o t o h p e c r o F r i A . S . U
Forensic anthropologist s working in a lab near Baghdad cata-
logued the injuries suf fered by every individual from the mass grave shown above. Some of this evidence w as used in the tr ial of Saddam Hussein and helped lead to his conviction. Af ter the trail, the human remains were turne d over to Kurdish officials for reburial y r o t a r o b a L n o i t a c i f i t n e d I A N D s e c r o F d e m r A , g n i K g i a r C
Upper Right: A forensic anthropologis t works in Vietnam
in 2006 as part of a militar y team, with assistance from local villagers, searching for remains of pilots shot down during the Vietnam War. Lower Right: Heather Th ew, who was trained as an anthropol-
ogist, is shown working at the Armed Forces DNA Laborator y where remains of missing soldiers are identified.
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Credits This constitutes an extension of the copyright page. We have made every ef fort to trace the ownership of all copyrighted material and to secure permission from copyright holders. In the event of any question arising as to the use of any material, we will be pleased to make the necessary corrections in future printings. Thanks are due to the following authors, publishers, and agents for permission to use the material indicated. 2, Cristina G. Mittermeier; 4, Courtesy, Peter Jones; 5, NASA; 6, (a) Lynn Kilgore, (b) NASA/
Space Telescope Science Institute, (c) Museum of Primitive Art and Culture, Peace Dale, RI, (e) Lynn Kilgore, (f) Justin Horocks/iStockphoto; 11, top, © Kenneth Garrett/NGS Image Collection; center, © Russell L. Ciochon, University of Iowa; bottom, Lynn Kilgore; 12, top, Courtesy, Judith Regensteiner; bottom, Courtesy, Kathleen Galvin; 13, top left, Robert Jurmain; top right, Courtesy, Nelson Ting; 14, top left and right, Lynn Kilgore; bottom, © Reuters/Corbis; 15, top, Courtesy, Linda Levitch; bottom, Courtesy, Julie Lesnik; 16, Nanette Barkey; 22, background, © Jamie VanBuskirk/istockphoto; bottom, © Diane France; 23, top and center left, U.S. Army Corps of Engineers, and the Regime Crime Liaison Office; center right, U.S. Air Force photo by Staff Sgt. Derrick C. Goode; bottom, Craig King, Armed Forces DNA Identification Laboratory; 24, Rolbos/iStockphoto; 27, Wikipedia; 28, J. van ( Johannes) Loon/Wikimedia Commons; 30, American Museum of Natural History; 31, Wikipedia; 32, top, With permission from the Master of Haileybury; bottom, © National Portrait Gallery, London; 33, top left and right, Lynn Kilgore; bottom, The Natural History Museum, London; 34, © Bettmann/Corbis; 35, © Gordon Chancellor; 36, bottom, Lynn Kilgore; 37, wolf: John Giustina/Getty Images; Dogs surrounding wolf: Lynn Kilgore and Lin Marshall; 38, © National Portrait Gallery, London; 39, © Russell L. Ciochon, University of Iowa; 40, (a) Michael Tweedie/Photo Researchers; (b) Breck P. Kent/ Animals Animals; 45, © Bettmann/Corbis; 48, Adrian T. Sumner/Photo Researchers, Inc.; 50, Courtesy, Dr. Michael S. Donnenberg; 51, top, Professors P. Motta and T. Naguro/SPL/ Photo Researchers, Inc.; bottom, A. Barrington Brown/Photo Researchers, Inc.; 58, Lynn Kilgore; 59, © Biophoto Ass ociates/Science Source/Photo Researchers; 67, Cellmark Diagnostics, Abingdon, UK; 70, Medi-Mation Ltd/Photo Researchers, Inc.; 72, (a) and (b) Wikipedia; 72, left, Raychel Ciemma and Precision Graphics; 79, Courtesy, Ray Carson, University of Florida News and Public Affairs; 80, (a–f) Lynn Kilgore; (g) Robert Jurmain; 86, Lynn Kilgore; 87, (a) and (b) © Dr. Stanley Flegler/Visuals Unlimited; 92, Louie Psihoyos/ Corbis; 105, (a) © John Cancalosi/Alamy; (b) © Phil Degginger/Alamy; (c) Elwyn Simons; (d) Marvin Dembinsky Photo Associates/Alamy; (e) Shoshannah White/Aurora; 106, Kazuo Unno/Minden Pictures; 107, left, Zhoukoudian Museum, China and © Russell L. Ciochon; right, © Russell L. Ciochon; 111, bottom left, J. C. Stevenson/Animals Animals; bottom right, Tom McHugh, Photo Researchers, Inc.; 113, © Corbis. All Rights Reserved; 116, © Dmitry Rukhlenko/iStockphoto; 119, (a–e), Lynn Kilgore; 121, Lynn Kilgore; 122, © Steve Geer/ iStockphoto; 124, Howler species: Raymond Mendez/Animals Animals; Spider monkey: Robert L. Lubeck/Animals Animals; Prince Bernhard’s titi: Marc van Roosmalen; Tamarin: © Zoological Society of San Diego, photo by Ron Garrison; Muriqui: Andrew Young; Whitefaced capuchins: © Jay Dickman/Corbis; Squirrel monkey: © Kevin Schafer/Corbis; Uakari: R. A. Mittermeier/Conservation International; 125, Baboon: Courtesy, Bonnie Pedersen/ Arlene Kruse; Macaque: Courtesy, Jean De Rousseau; Gibbon: Lynn Kilgore; Tarsier: Courtesy, David Haring, Duke University Primate Zoo; Orangutan: © Tom McHugh/Photo Researchers, Inc.; Langur: Joe MacDonald/Animals Animals; Lemur: Courtesy, Fred Jacobs; Loris: Courtesy, San Francisco Zoo; Cercopithecus: Robert Jurmain; Colobus: Robert Jurmain; Galagos: Courtesy, Bonnie Pedersen/Arlene Kruse; Chimpanzee: Courtesy, Arlene Kruse/Bonnie Pedersen; Mountain gorilla: Lynn Kilgore;126, © Russell L. Ciochon, University of Iowa; 127, Redrawn by Stephen Nash from original art in John G. Fleagle, Primate Adaptation and Evolution, 2nd ed., 1999. Reprinted by permission of publisher and Stephen Nash.; 128, Lynn Kilgore; 131, top left and right, Lynn Kilgore; center, © Viktor Deak, after John G. Fleagle; 132, top left and right, Courtesy, Fred Jacobs; bottom left, Courtesy, San Francisco Zoo; bottom right, Courtesy, Bonnie Pedersen/Arlene Kruse; 133, David Haring, Duke University Primate Zoo; 134, top left, © Andrew Young; top right, © Kevin Schafer/Corbis; center right, Marc van Roosmalen; bottom right, R. A. Mittermeier/ Conservation International; bottom left, © Jay Dickman/Corbis; 135, top left, © Zoological Society of San Diego, photo by Ron Garrison; top right, Raymond Mendez/Animals Animals; bottom, Robert L. Lubeck/Animals Animals; 136, Robert Jurmain; 137, top left and right, Courtesy, Bonnie Pedersen/Arlene Kruse; bottom, Courtesy, Nelson Ting; 139, top, Lynn Kilgore; bottom left, Noel Rowe; bottom right, Lynn Kilgore; 140, bottom left, right, Lynn Kilgore; 141, (a) and (b), Lynn Kilgore; 142, (a) and (b), Lynn Kilgore; 144, Courtesy, Ellen Ingmanson; 145, Lynn Kilgore; 147, John Oates; 148, Karl Ammann; 149, WildlifeDirect.org; 152, © Doug Berry/iStockphoto; 155, Courtesy, Jean De Rousseau; 156, top, Russ Mittermeir; bottom, Lynn Kilgore; 158, Time Life Pictures/Getty Images; 160, top and bottom, Lynn Kilgore; 162, Curt Busse; 163, top and bottom, Lynn Kilgore; 165, top left and bottom right, Courtesy, Arlene Kruse/Bonnie Pedersen; top right, Courtesy, Meredith Small; bottom left, Robert Jurmain; 166, Ireven De Vore/Anthro-Photo; 167, whozoo.org; 168, Joe MacDonald/Animals Animals; 169, © Peter Henzi; 170, top left, Courtesy, David Haring, Duke University Primate Center; top right, Courtesy, Arlene Kruse/Bonnie Pedersen; center left, Robert Jurmain; center right, Robert Jurmain; bottom, © Tom McHugh/Photo Researchers, Inc.; 171, top, Harlow Primate L aboratory, University of Wisconsin; bottom, Lynn Kilgore; 172, left, Lynn Kilgore; right, Manoj Shah/The Image Bank; 173, © Thomas Breuer—WCS; 174, Courtesy, Tetsuro Matsuzawa; 175, top, Barth Wright/EthoCebus Project; bottom, Noemi Spagnoletti/EthoCebus Project; 176, Lynn Kilgore; 178, Rose A. Sevcik, Language Research Center, Georgia State University; photo by Elizabeth Pugh; 179, (a) Lynn Kilgore; (b) Allposters.com; 182, Robert Jurmain; 185, top, Wikipedia; bottom,
Russell L. Ciochon; 188, top, Natural History Museum, London; bottom, David Pilbeam; 194, Russell L. Ciochon; 198, top, Barry Le wis; bottom, John Reader/Photo Researchers, Inc.; 202, Toros-Menalla: © Mission Paléoanthropologique Franco-Tchadienne; W T 17000: Reproduced with permission of the National Museums of Kenya, copyright reserved, courtesy Alan Walker; Lucy: © Institute of Human Origins; SK 48: © Russell L. Ciochon, University of Iowa; Sts 5: © Russell L . Ciochon, University of Iowa; Taung Child: Courtesy, P. V. Tobias, photo by Alun Hughes; “Zinj”: © Jeffrey Schwartz, University of Pittsburg; 203, John Hodgkiss; 204, © Mission Paléoanthropologique Franco-Tchadienne; 205, © David L. Brill/Atlanta; 207, Lynn Kilgore; 208, Courtesy, Peter Jones; 209, top, Institute of Human Origins; bottom left, © Russell L. Ciochon, University of Iowa; bottom right, Carol Ward, University of Missouri; 210, © Zeresenay Alemseged, Max Planck Institute for Evolutionary Anthropology, Leipzig; 212, ER 406: Reproduced with permission of the National Museums of Kenya, copyright reserved; WT 17000: Reproduced with permission of the National Museums of Kenya, copyright reserved, courtesy Alan Walker; SK 48: © Russell L. Ciochon, University of Iowa; ER 729: Reproduced with permission of the National Museums of Kenya, copyright reserved; ER 732: Reproduced with permission of the National Museums of Kenya, copyright reserved; OH 5 “Zinj”: © Jeffrey Schwartz, University of Pittsburg; 213, © Russell L. Ciochon, University of Iowa; bottom left and right, Alun Hughes, reproduced by permission of Professor P. V. Tobias; 214, © Russell L. Ciochon, University of Iowa; 216, left and right, Reproduced with permission of the National Museums of Kenya, copyright reserved; 217, ER 1813, ER 1470: Reproduced with permission of the National Museums of Kenya, copyright reserved; AL 666-1: © William H. Kimbel, Institute Human Origins; Stw 53: Courtesy, P. V. Tobias (reconstruction by Ronald J. Clarke); OH 7: © Russell L. Ciochon, University of Iowa; 222, © Russell L. Ciochon, University of Iowa; 223, top, Institute of Human Origins, photo by Nanci Kahn; top middle, bottom middle, and bottom, and right, Institute of Human Origins, photo by Don Johanson; 222–223, background, © Britta Kasholm-Tengve/iStockphoto; 224, David Lordkipanidze; 226, Ceprano: © Giorgio Manzi, Universitá di Roma; Dmanisi: Courtesy, David Lordkipanidze; East Turkana, Olduvai: © Jeffrey Schwartz, University of Pittsburg; West Turkana: Reproduced with permission of the National Museums of Kenya, copyright reserved; 227, Zhoukoudian: © Russell L. Ciochon, University of Iowa; Lantian, Hexian, Ngandong: Courtesy, Milford Wolpoff; Trinii: Courtesy, S. Sartano; Sangiran: © Russell L. Ciochon, University of Iowa; 230, Zhoukoudian: © Russell L. Ciochon, University of Iowa; Sangiran 17: Courtesy, Milford Wolpoff; Ngandong 5: © Russell L. Ciochon, University of Iowa; Zhoukoudian: © Russell L. Ciochon, University of Iowa; ER 3733: Courtesy, Milford Wolpoff; O.H. 9: © Jeffrey Schwartz, University of Pittsburg; 231, © Jeffrey Schwartz, University of Pittsburg; 232, Reproduced with permission of the National Museums of Kenya, copyright reserved; 338, (a, b, and c) David Lordkipanidze; 234, David Lordkipanidze; 235, S. Sartano; 236, top, bottom, © Russell L. Ciochon, University of Iowa; 238, (a) and (b) Courtesy, Milford Wolpoff; 240, © Giorgio Manzi, Universitá di Roma; 241, The Museum of Primitive Art and Culture, Peace Dale, RI, photo by William Turnbaugh; 246, Javier Trueba/Madrid Scientific Films/Photo Researchers; 250, top, Courtesy, Milford Wolpoff; bottom, © Robert Franciscus, University of Iowa; 251, Courtesy, Milford Wolpoff; 252, Arago: Courtesy, H. DeLumley; Steinheim: Courtesy, Milford Wolpoff; Petralona: Courtesy, Milford Wolpoff; Bodo: © Robert Franciscus, University of Iowa; Florisbad: Courtesy, Günter Bräuer; Kabwe: Courtesy, Milford Wolpoff; 253, Jinniushan: © Russell L. Ciochon, University of Iowa; Dali: Courtesy, Milford Wolpoff; 255, top, Courtesy, Milford Wolpoff; bottom, © Russell L. Ciochon, University of Iowa; 260, Shanidar I: Courtesy, Erik Trinkaus; Gibraltar: © Robert Franciscus, University of Iowa; Amud 1: Courtesy, Milford Wolpoff; La Chapelle: Courtesy, Fred Smith; St. Césaire: Harry Nelson; La Ferrassie 1: Courtesy, Fred Smith; 261, St. Césaire: Harry Nelson; La Ferrassie: Courtesy, Fred Smith; Krapina/Vindija: Courtesy, Milford Wolpoff; Amud: Courtesy, Milford Wolpoff; Shanidar: Erik Trinkaus, Washington University, St. Louis; Tabun/Kebara: Courtesy, Fred Smith; La Chapelle: Courtesy, Fred Smith; Gibraltar: © Robert Franciscus, University of Iowa; 262, top, Courtesy, Fred Smith; bottom, Harry Nelson; 263, (a) and (b), Courtesy, Fred Smith; 264, bottom left, Harry Nelson; bottom right, Courtesy, Erik Trinkaus, Washington University, St. Louis; 267, © Randall White, New York University; 276, The Gallery Collection/Corbis; 280, Skhu¯ l, Courtesy, Fred Smith; Jebel Qaf zeh: Courtes y, Milford Wolpoff; Herto: David L . Brill/ Atlanta; Border Cave: Courtesy, Fred Smith; Klasies River Mouth, Courtesy, Fred Smith; Omo: Courtesy, Milford Wolpoff; 283, Russell L. Ciochon; 283, Jebel Qafzeh 6: Courtesy, Milford Wolpoff ; Border Cave 1: Courte sy, Fred Smith; Skhu¯l 5: Courte sy, Fred Smith; Prˇedmostí; Harry Nels on; Cro-Magnon I: Courtesy, David Frayer; 284, David L. Brill/Atlanta; 285, top, Courtesy, David Frayer; bottom left, Courtesy, David Frayer; bottom right, Milford Wolpoff; 288, Zhoukoudian: © Russell L. Ciochon, University of Iowa; Lake Mungo: Courtesy, Milford Wolpoff; Kow Swamp: Courtesy, Milford Wolpoff; 289, © Mircea Gerhase; 290, Combe Capelle: © Staatliche Museen zu Berlin, Museum fur Vor- und Fruhgeschichte; Mladecˇ: Courtesy, Milford Wolpoff ; Oase: © Erik Trinkaus , Washington Univer sity, St. Louis; Cro-Magnon: Courtesy, David Frayer; 291, (a), Milford Wolpoff; (b), © Robert Franciscus; 292, (a), Courtesy, David Frayer; (b), Courtesy, Milford Wolpoff; 294, © Peter Brown; 295, N. O. Bader; 299, Altamira: Reproduced from Emile Cartailhac and l’abbé Henri Breuil (1906); Brassempouy: Harry Nelson; Sungir: © Randall White, New York University; Katanda: Courtesy, Chip Clark , Smithsonian Institution; Blombos Cave: © CS Henshilwood, Iziko Museums of Cape Town; Apollo 11 Cave: © Gerald Newlands, University of Calgary ; 302, Jose Fuste Raga/Corbis; 307, (a), © Peter Johnson/Corbis; (b), © Charles & Josette Lenars/ Corbis; (c), © Gallo Images/Corbis; (d), © Otto L ang/Corbis; (e), Lynn Kilgore; 308, © Tannen Maury/epa/Corbis; 309, (a) and (b), Robert Jurmain; 314, © Atlantide Phototravel/ Corbis; 315, © Michael S . Yamashita/Corbis; 319, Reuters/STR/Landov; 320, © Biophoto Associates/Photo Researchers, Inc.; 323, (a) Renee Lynn/Photo Researchers; (b) George Holton/Photo Researchers; 325, (a) © Rafal Belzowski/iStockphoto; (b) © Danny Warren/ iStockphoto; 328, CDC/Jean Roy; 329, Karl Ammann; 330, Hoang Dinh Nam/AFP/Getty Images; 334, left, Lynn Kilgore; center, Robert Jurmain; right, Lynn Kilgore; 335, top right and top left, Robert Jurmain; center, Charlotte Roberts; bottom, Courtesy, Art Aufderheide; 336, © Podvysotskiy Roman/iStockphoto; 341, Charlotte Rogers; 351, James F. O’Connell; 355, left, © Steve Bloom/stevebloom.com; right, © Karl Ammann/Corbis; 358, Terrance Emerson/iStockphoto; 362, top, Lynn Kilgore; bottom left, Brasil2/iStockphoto; bottom right, © Russ Mittermeier; 364, Robert Jurmain; 365, top, Wilbur Garrett/National Geographic Society/Getty Images; bottom, Louis Meulstee/Peter Arnold, Inc.; 367, Lynn Kilgore; 369, Photo of earth: Earth Data Analysis Center; 374, (a) © Klaus Meindl/iStockphoto; (b) © A. Goodreds/iStockphoto; (c) © Glenn Rogers/iStockphoto; (d) © Dave Logan/iStockphoto; (e) © Tyler Oliver/iStockphoto
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