I. Author Bio

II. Author Letter

I. Author Bio

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Craig Stanford is a professor of anthropology and biological sciences at the University of Southern California, where he also directs the Jane Goodall Research Center. He has conducted field research on primate behavior in south Asia, Latin America, and East Africa. He is well known for his long-term studies of meat-eating among wild chimpanzees in Gombe, Tanzania, and of the relationship between mountain gorillas and chimpanzees in the Impenetrable Forest of Uganda. He has authored or coauthored more than 120 scientific publications. Craig has received USC’s highest teaching awards for his introductory biological anthropology course. In addition, he has published eleven books on primate behavior and human origins, including Significant Others (2001), Upright (2003) and Beautiful Minds (2008). He and his wife, Erin Moore, a cultural anthropologist at USC, live in South Pasadena, California, and have three children.

John Allen is a research scientist in the Dornsife Cognitive Neuroscience Imaging Center and the Brain and Creativity Institute at the University of Southern California. Previously, he was a neuroscience researcher at the University of Iowa College of Medicine and a faculty member in the Department of Anthropology at the University of Auckland, New Zealand, for several years. His primary research interests are the evolution of the human brain and behavior, and behavioral disease. He also has research experience in molecular genetics, nutritional anthropology, and the history of anthropology. He has conducted fieldwork in Japan, New Zealand, Papua New Guinea, and Palau. He has received university awards for teaching introductory courses in biological anthropology both as a graduate student instructor at the University of California and as a faculty member at the University of Auckland. In addition to BiologicalAnthropology, he is also the author of Medical Anthropology: A Biocultural Approach (with Andrea S. Wiley; 2009) and The Lives of the Brain (2009). John and his wife, Stephanie Sheffield, have two sons, Reid and Perry.

Susan Antón is a professor in the Center for the Study of Human Origins, Department of Anthropology at New York University, where she also directs the M.A. program in Human Skeletal Biology. Her field research concerns the evolution of genus Homo in Indonesia and human impact on island ecosystems in the South Pacific. She is best known for her work on H. erectus in Kenya and Indonesia, for which she was elected as a fellow of the American Association for the Advancement of Science (AAAS) in 2008. She is past editor of the Journal of Human Evolution. She received awards for teaching as a graduate student instructor of introductory physical anthropology and anatomy at the University of California, was Teacher of the Year while at the University of Florida, and a Golden Dozen teaching award recipient at NYU. She has been twice elected to Who’s Who Among America’s Teachers. Susan and her husband, Carl Swisher, a geochronologist, raise Anatolian shepherd dogs.

Dear Colleague,

It is our pleasure to be able to bring you the third edition of our textbook Biological anthropology: the natural history of

humankind.We are writing to you to share some highlights from the new edition. We have done our best to keep the book comprehensive, cutting edge, and accessibly readable. Over the past three years, new fossil discoveries, new revelations about primate behavior, and new breakthroughs in molecular biology have made an update of our previous edition essential. As always, we have endeavored to provide students and instructors with the very best coverage of these issues, and also the best photographs and images available.

We believe that Biological anthropology offers you an outstanding choice in its comprehensive coverage of topics as well

as its clarity, originality, critical-thinking approach, and presentation of beautifully done artwork and photography. All the traditional topics covered in other introductory biological anthropology texts are covered in detail. We also treat topics that are rarely covered in traditional texts, but are at the cutting edge of field. These include biomedical anthropology, brain evolution, and forager societies.

In addition, new aspects of the third edition include:

• Updated treatment of recent discoveries of Australopithecus sediba, Ardipithecus ramidus, Darwinius, and Denisova hominins,

plus advances in the study of Ancient DNA of Neandertals

• New discoveries about chimpanzee culture, including the latest research on tool use and hunting

• Updates on genetics, including ancient DNA and population genetics

• Expanded treatment of evolutionary aspects of human health and disease

• New photographs of fossils, primates, and other subjects

• New anatomical illustrations, featuring systematic redrawing of fossil and osteological artwork by medical artists

Each chapter of the book now contains an updated visual summary. Each chapter also features links, where appropriate, to

MyAnthroLab, which provides highly useful information and exercises for use in labs that accompany a biological anthropology course. MyAnthroLab is available at no cost to students who are using Biological anthropology.

We wrote the first edition of Biological anthropology because, as teaching assistants during our graduate school days, we

were disappointed in the overall quality of other books that were available. We are very pleased that Biological anthropology

has become a leader in the biological anthropology textbook market. We are, as always, grateful for feedback from instructors and we try our best to incorporate updates to both the content and appearance of the book with instructor needs in mind. We realize that you, the instructor, have a choice of books to assign and we were committed to producing a new third edition that would meet your needs and those of students in this fascinating field of study.

Sincerely yours,

Craig Stanford

John S. Allen

Susan c. Antón

Introduction: What Is Biological Anthropology? 

Part I: Mechanisms of Evolution
Chapter 1: Origins of Evolutionary Thought   
Chapter 2: Genetics: Cells and Molecules   
Chapter 3: Genetics: From Genotype to Phenotype   
Chapter 4: The Forces of Evolution and the Formation of Species   
Chapter 5: Human Variation: Evolution, Adaptation, and Adaptability  

Part II: Primates
Chapter 6: The Primates   
Chapter 7: Primate Behavior  

Part III: Paleontology and Primate Evolution
Chapter 8: Fossils in Geological Context   
Chapter 9: Origin of Primates   
Chapter 10: Becoming Human: The Ape—Hominin Transition  

Part IV: The Human Fossil Record
Chapter 11: Early Hominins   
Chapter 12: Rise of the Genus Homo   
Chapter 13: Archaic Homo sapiens and Neandertals   
Chapter 14: The Emergence and Dispersal of Homo sapiens  

Part V: New Frontiers in Biological Anthropology
Chapter 15: Evolution of the Brain and Language   
Chapter 16: Biomedical Anthropology   
Chapter 17: The Evolution of Human Behavior   
Chapter 18: Bioarchaeology and Forensic Anthropology  

Appendix A: Overview of the Brain  
Appendix B: Primate and Human Comparative Anatomy  
Appendix C: The Hardy—Weinberg Equilibrium  
Appendix D: Metric—Imperial Conversions  

COMPREHENSIVE TABLE OF CONTENTS:

Preface 
About the Authors 

Introduction: What Is Biological Anthropology? 

The Scope of Biological Anthropology 
Paleoanthropology 
Skeletal Biology and Human Osteology 
Paleopathology 
Forensic Anthropology 
Primatology 
Human Biology 
The Roots of Modern Biological Anthropology 
Anthropology and Its Subfields 
Cultural Anthropology 
INSIGHTS AND ADVANCES: A Paradigm Split in Anthropology? 
Archaeology 
Linguistic Anthropology 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Part I: Mechanisms of Evolution

Chapter 1: Origins of Evolutionary Thought 
What Is Science? 
The Early Thinkers 
The Roots of Modern Science 
Linnaeus and the Natural Scheme of Life 
The Road to the Darwinian Revolution 
The Uniformitarians: Hutton and Lyell 
The Darwinian Revolution 
The Galápagos 
Refining the Theory of Evolution by Natural Selection 
INSIGHTS AND ADVANCES: Darwin versus Wallace? 
The Response to Darwin 
Science and Creationism 
INSIGHTS AND ADVANCES: What Is Intelligent Design? 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 2: Genetics: Cells and Molecules 
Genetics 
The Study of Genetics 
Genetic Metaphors: Blueprints, Recipes, or What? 
The Cell 
Cell Anatomy  
INSIGHTS AND ADVANCES: Cloning Controversies 
DNA Structure and Function 
DNA Structure I: The Molecular Level 
DNA Function I: Replication 
DNA Function II: Protein Synthesis 
DNA Structure II: Chromosomes and Cell Division 
INNOVATIONS: The Wide World of RNA 
INSIGHTS AND ADVANCES: Biochemical Individuality 
Molecular Tools For Bioanthropological Research 
Indirect versus Direct Research Methods 
PCR, Mitochondrial DNA, and Ancient DNA 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 3: Genetics: From Genotype to Phenotype 
From Genotype to Phenotype 
The ABO Blood Type System 
Obesity: A Complex Interaction 
Mendelian Genetics 
Mendel’s Postulates 
Linkage and Crossing Over 
Mutation 
Point Mutation and Sickle Cell Disease 
Trinucleotide Repeat Diseases 
Mutations: Bad, Neutral, and Good 
X-Linked Disorders 
Mendelian Genetics in Humans 
Genetics Beyond Mendel  
INSIGHTS AND ADVANCES: Popular Mendelism and the Shadow of Eugenics 
Polygenic Traits, the Phenotype, and the Environment 
Heritability and IQ Test Score Performance 
Phenylketonuria: Illustrating Mendelian and Post-Mendelian Concepts 
Genes and Environments 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 4: The Forces of Evolution and the Formation of Species 
How Evolution Works 
Where Does Variation Come From? 
How Natural Selection Works 
Other Ways by Which Evolution Happens 
Classification and Evolution 
Taxonomy and Speciation 
What Is a Species? 
A Guide to Species Concepts 
Reproductive Isolating Mechanisms 
The Origin of Species: How Species Are Formed 
INSIGHTS AND ADVANCES: What’s in a Name? Species Concepts, Genetics, and Con-servation 
The Tempo of Speciation 
Adaptation 
Is Everything Adaptive? 
Hardy—Weinberg Equilibrium 
Levels of Selection 
Inclusive Fitness 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 5: Human Variation: Evolution,  Adaptation, and Adaptability 
Human Variation at the Individual and Group Level 
What Is a Population? 
Historical Perspectives on Human Variation 
Recording Human Variation in Past Civilizations 
The Monogenism—Polygenism Debate 
Race and Racism in the Twentieth Century 
Changing Attitudes toward Race in Anthropology 
Deconstructing Racial Features 
Population Genetics 
Polymorphisms: ABO and Other Blood Type Systems 
Gene Flow and Protein Polymorphisms 
Polymorphisms and Phylogenetic Studies 
Polymorphisms and Natural Selection in Human Populations 
The Evolution of Lactose Tolerance 
Balanced Polymorphisms: Sickle-Cell and Other Conditions 
Adaptation and Adaptability 
Levels of Adaptability 
Heat and Cold  
INSIGHTS AND ADVANCES: Technology and Extreme Environments 
Body Size and Shape 
Living at High Altitude 
Skin Color 
Adaptability to Water 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Part II: Primates

Chapter 6: The Primates 
The Primate Radiation 
The Extraordinary Diversity of Nonhuman Primates 
What Exactly Is a Primate? 
Anatomical Traits 
Life History Traits 
Behavioral Traits: Activity and Sociality 
A Guide to the Nonhuman Primates 
The Strepsirhines 
The Haplorhines  
INSIGHTS AND ADVANCES: The Rarest of the Rare 
The New World Monkeys 
The Old World Monkeys 
The Hominoids 
INSIGHTS AND ADVANCES: The Impending Extinction of the Great Apes? 
Primate Ecology 
The Cycles of a Tropical Forest 
You Are What You Eat: Dietary and Digestive Strategies 
Diet and Feeding Competition 
Territories and Ranges 
Predation 
Primate Communities 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 7: Primate Behavior 
Studying Primates 
The Evolution of Primate Social Behavior 
Social Behavior and Reproductive Asymmetry 
Male Reproductive Strategies 
Female Reproductive Strategies 
Why Are Nonhuman Primates Social? 
The Paradox of Sociality 
INNOVATIONS: Culture in Nonhuman Primates 
Types of Nonhuman Primate Societies 
INSIGHTS AND ADVANCES: The Infanticide Wars 
INSIGHTS AND ADVANCES: Are Chimpanzees from Mars and Bonobos from Venus? 
Reconstructing the Evolution of Primate Societies 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Part III: Paleontology and Primate Evolution

Chapter 8: Fossils in Geological Context 
How to Become a Fossil 
The Importance of Context 
Stratigraphy 
The Geologic Time Scale 
How Old Is It? 
Relative Dating Techniques 
Calibrated Relative Dating Techniques 
INSIGHTS AND ADVANCES: The Piltdown Hoax 
Chronometric Dating Techniques 
INSIGHTS AND ADVANCES: Dating Controversies 
INNOVATIONS: Time in a Bottle 
The Earth in the Cenozoic 
Continents and Land Masses 
The Environment in the Cenozoic 
Overview of Climatic Changes during the Cenozoic 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 9: Origin of Primates 
The Mesozoic and Beyond 
Dawn of the Age of Mammals 
The Crater of Doom: What Happened at the K—T Boundary? 
Changes in the Paleocene: The Origin of Primates? 
Why Primates? 
Early Primates of the Eocene 
Adapoids (Strepsirhine Ancestors) 
Omomyoids (Haplorhine Ancestors) 
Continental Drift and Eocene Primates 
INSIGHTS AND ADVANCES: Subfossil Lemurs of Madagascar 
Selective Pressures Favoring the Strepsirhine—Haplorhine Split 
Evolution of Higher Primates 
The First Monkeys? 
New World Monkeys 
Old World Monkeys 
What Favored the Origin of Anthropoids? 
The Earliest Apes 
Selection Pressures and the Divergence of Monkeys and Apes 
The Monkey’s Tale: What Happened to Primate Diversity in the Miocene? 
Molecular Evolution in Primates 
A Primate Molecular Phylogeny 
Molecular Phylogeny and Human Origins 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 10: Becoming Human: The Ape—Hominin Transition 
Becoming a Biped 
Anatomical Changes 
Constructing the Bipedal Body Plan 
Locomotion of the Last Common Ancestor 
Why Bipeds? 
INSIGHTS AND ADVANCES: Overheated Radiator? 
The Transition to Human Behavior 
Primate Intelligence: Why Are Human Brains Big? 
What Made Humans Human? 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Part IV: The Human Fossil Record

Chapter 11: Early Hominins 
Will You Know a Hominin When You See One? 
INSIGHTS AND ADVANCES: A Rose by Any Other Name: Hominins versus Hominins 
The First Hominins? 
Sahelanthropus tchadensis 
Orrorin tugenensis 
Ardipithecus ramidus and Ardipithecus kadabba 
INSIGHTS AND ADVANCES: Treasures of the Afar Triangle 
Australopithecus and Kin 
Australopithecus anamensis 
Australopithecus afarensis 
INNOVATIONS: Dikika and Development 
Australopithecus bahrelghazali 
Kenyanthropus platyops 
Australopithecus garhi 
Australopithecus africanus 
The Robust Australopithecines (or Paranthropines) 
Understanding the Australopithecine Radiation 
Cohabitation 
Tools and Intelligence 
Ancestors and Descendants 
Questions for Future Paleoanthropologists 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 12: Rise of The Genus Homo 
Defining the Genus Homo 
Earliest Genus Homo 
Early Tool Use 
Hunting and Scavenging 
INSIGHTS AND ADVANCES: Understanding the Meat-Eating Past through the Pre-sent 
Who Was Homo erectus? 
Anatomical Features 
Homo erectus versus Homo ergaster 
Homo erectus around the World 
African Origins 
The First African Diaspora: Republic of Georgia 
Dispersal into East Asia 
The Status of Homo erectus in Europe 
The Lifeways of Homo erectus 
Homo erectus and the Early Stone Age 
A Higher-Quality Diet: Homo erectus Subsistence 
Homo erectus Life History 
INSIGHTS AND ADVANCES: What’s Size Got to Do with It? 
Homo erectus Leaves Africa 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 13: Archaic Homo sapiens and Neandertals 
Hominin Evolution in the Middle to Late Pleistocene 
Defining Anatomically Modern Homo sapiens 
Archaic Homo sapiens 
European Archaic Homo sapiens 
African Archaic Homo sapiens 
Asian Archaic Homo sapiens 
Behavior of Archaic Homo sapiens 
Stone Tools 
Biodegradable Tools 
Big Game Hunting 
Fire, Campsites, and Home Sites 
The Neandertals 
Geographic and Temporal Distribution 
History of Neandertal Discovery 
INSIGHTS AND ADVANCES: Neandertal Image Makeovers 
Neandertal Anatomy and DNA 
Growth and Development 
Health and Disease 
INNOVATIONS: Neandertal Genes 
Neandertal Behavior 
Material Culture 
Coping with Cold 
Hunting and Subsistence 
Cannibalism 
Burials 
Ritual and Symbolic Behavior 
Phylogenetic and Taxonomic Issues: An Overview 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 14: The Emergence and Dispersal of Homo sapiens 
The Emergence of Modern Humans 
Models of Modern Human Origins 
Multiregional and Replacement Models 
Predictions of the Two Models 
Anatomy and Distribution of Early Humans 
Africa 
Near East 
Europe 
Asia and Southeast Asia 
Australia 
INSIGHTS AND ADVANCES: The Little People of Flores 
Archaeology of Modern Human Origins 
Stone and Other Tools 
Subsistence 
Symbolism 
Settlement of the New World and Pacific Islands 
Molecular Genetics and Human Origins 
INSIGHTS AND ADVANCES: The “Vitamin D Line” 
Mitochondrial DNA 
The Y Chromosome 
MRCAs for Nuclear Genes 
Ancient DNA 
Interpreting Models of Human Origins 
Paleontology and Archaeology 
Molecular Genetics 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Part V: New Frontiers in Biological Anthropology

Chapter 15: Evolution of the Brain and Language 
Issues in Hominin Brain Evolution 
Brain Size and Encephalization 
Brain Size and the Fossil Record 
Brain Reorganization 
INSIGHTS AND ADVANCES: The Ten-Percent Myth: Evolution and Energy 
Language: Biology and Evolution 
The Evolution of Grammar 
Language in the Brain 
Language in the Throat 
Language Ability and the Fossil Record 
INNOVATIONS: Music, the Brain, and Evolution 
INSIGHTS AND ADVANCES: Ape Language Studies 
Scenarios of Language Evolution 
Brain Size, Language, and Intelligence 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 16: Biomedical Anthropology 
Epidemiology: Basic Tools For Biomedical Anthropology 
Rates: Mortality, Incidence, and Prevalence 
Epidemiological Transitions 
Biocultural and Evolutionary Approaches to Disease 
The Biocultural Approach 
The Evolutionary Approach 
Birth, Growth, and Aging 
Human Childbirth 
Patterns of Human Growth 
Stages of Human Growth 
The Secular Trend in Growth 
Menarche and Menopause 
Aging 
Infectious Disease and Biocultural Evolution 
Human Behavior and the Spread of Infectious Disease 
INSIGHTS AND ADVANCES: Kuru, Cannibalism, and Prion Diseases 
Infectious Disease and the Evolutionary Arms Race 
Diet and Disease 
The Paleolithic Diet 
Agriculture and Nutritional Deficiency 
Agriculture and Abundance: Thrifty and Nonthrifty Genotypes 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 17: The Evolution of Human Behavior 
Studying the Evolution of Human Behavior 
The Evolution of Human Behavior: Four Approaches 
Behavioral Patterns and Evolution 
Traditional Lives in Evolutionary Ecological Perspective 
Quantification in Evolutionary Ecological Research 
Hunting, Gathering, and the Sexual Division of Labor 
Sexual Selection and Human Behavior 
Risk-Taking Behavior 
Inbreeding Avoidance and Incest Taboos 
Language-Related Cross-Cultural Behaviors 
Motherese or Infant-Directed Speech 
Basic Color Terms 
INSIGHTS AND ADVANCES: Reading, Writing, and Evolution 
Behavioral Disease 
Depression and Natural Selection 
Schizophrenia 
Psychoactive Substance Use and Abuse 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

Chapter 18: Bioarchaeology and Forensic Anthropology 
Life, Death, and the Skeleton 
Field Recovery Methods 
Laboratory Processing, Curation, and Chain of Custody 
The Biological Profile 
Age at Death 
Sex 
Ancestry 
Height and Weight 
INSIGHTS AND ADVANCES: Ancestry Genetics 
Premortem Injury and Disease 
Taphonomy 
Perimortem Trauma 
Postmortem Trauma 
DNA, Kinship, and Identity 
Identification and Forensic Anthropology 
INSIGHTS AND ADVANCES: The Genghis Khan Effect 
Time Since Death 
Antemortem Records and Positive IDs 
Facial Reconstruction 
Applications of Bioarchaeology 
Mortuary Archaeology 
Biocultural Evolution of Health and Disease 
Activity Patterns and Subsistence Change 
Applications of Forensic Anthropology 
Mass Fatalities 
War Dead 
War Crimes and Genocide 
Epilogue 
Summary • Critical Thinking Questions • Key Terms • Suggested Reading

APPENDIX A  Overview of the Brain 
APPENDIX B  Primate and Human Comparative Anatomy 
APPENDIX C  The Hardy-Weinberg Equilibrium 
APPENDIX D  Metric-Imperial Conversions 

Glossary 
Bibliography 
Photo Credits 
Index 

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