Exploring Biological Anthropology The Essentials I RB SE A MONT. KOS. S S The Modern World Australia and possessions China Denmark and possessions France and possessions India and possessions Italy Japan and possessions Netherlands and possessions New Zealand Norway and possessions Portugal and possessions Russian Federation Spain and possessions Turkey United Kingdom and possessions United States and possessions B-H = Bosnia-Herzegovina BI A SOUTH SUDAN This page intentionally left blank Exploring Biological Anthropology The Essentials Fourth Edition Craig Stanford University of Southern California John S. Allen University of Southern California Susan C. Antón New York University Boston Columbus Indianapolis New York San Francisco Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montréal Toronto Delhi Mexico City São Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo VP, Product Development: Dickson Musslewhite Publisher: Charlyce Jones-Owen Editorial Assistant: Laura Hernandez Program Team Lead: Maureen Richardson Project Team Lead: Melissa Feimer Program Manager: Rob DeGeorge Project Manager: Cheryl Keenan Art Director: Maria Lange Cover Art: Craig Stanford Director, Digital Studio: Sacha Laustein Digital Media Project Manager: Amanda A. Smith Procurement Manager: Mary Fischer Procurement Specialist: Mary Ann Gloriande Full-Service Project Management and Composition: Lumina Datamatics, Inc./Nancy Kincaid Printer/Binder: Courier/Kendallville Cover Printer: Phoenix Color/Hagerstown Text Font: SabonLTStd 10.5/12 Acknowledgements of third party content appear on page 482 which constitutes an extension of this copyright page. DK Maps designed and produced by DK Education, a division of Dorling Kindersley Limited, 80 Strand, London WC2R ORL. DK and the DK logo are registered trademarks of Dorling Kindersley Limited. Copyright © 2017, 2013, 2010 by Pearson Education, Inc. or its affiliates. All Rights Reserved. Printed in the United States of America. This publication is protected by copyright, and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise. For information regarding permissions, request forms and the appropriate contacts within the Pearson Education Global Rights & Permissions department, please visit www.pearsoned.com/permissions/. PEARSON, ALWAYS LEARNING, and REVEL are exclusive trademarks in the U.S. and/or other countries owned by Pearson Education, Inc. or its affiliates. Unless otherwise indicated herein, any third-party trademarks that may appear in this work are the property of their respective owners and any references to third-party trademarks, logos or other trade dress are for demonstrative or descriptive purposes only. Such references are not intended to imply any sponsorship, endorsement, authorization, or promotion of Pearson’s products by the owners of such marks, or any relationship between the owner and Pearson Education, Inc. or its affiliates, authors, licensees or distributors. Library of Congress Cataloging-in-Publication Data Stanford, Craig B. (Craig Britton) Exploring biological anthropology / Craig Stanford, University of Southern California, John S. Allen, University of Southern California, Susan C. Anton New York University.—Fourth edition.   pages cm ISBN 978-0-13-401401-2 (pbk.) 1. Physical anthropology—Textbooks. I. Allen, John S. (John Scott) II. Antón, Susan C. III. Title. GN25.S74 2017 599.9—dc23 2015035818 10 9 8 7 6 5 4 3 2 1 Student: ISBN-10: 0-13-401401-4 ISBN-13: 978-0-13-401401-2 A la Carte ISBN-10: 0-13-432383-1 ISBN-13: 978-0-13-432383-1 To Our Parents This page intentionally left blank Brief Contents Part I Foundations 1 What Is Biological Anthropology? 1 2 Origins of Evolutionary Thought 12 Part II Mechanisms of Evolution 3 Genetics: Cells and Molecules 4 Genetics: From Genotype to Phenotype 5 The Forces of Evolution and the Formation of Species 6 Human Variation: Evolution, Adaptation, and Adaptability Part III 59 83 105 Primates 7 The Primates 8 Primate Behavior Part IV 9 30 141 178 The Fossil Record Geology and Primate Origins 200 10 Early Hominins and Australopithecus 11 Rise of the Genus Homo 12 Archaic Homo sapiens and Neandertals 309 13 The Emergence, Dispersal, and Bioarchaeology of Homo sapiens Part V 242 277 342 Biology and Behavior of Modern Humans 14 Evolution of the Brain and Behavior 373 15 Biomedical and Forensic Anthropology 402 Appendix A Primate and Human Comparative Anatomy Appendix B The Hardy–Weinberg Equilibrium Appendix C Metric–Imperial Conversions 441 446 450 ix This page intentionally left blank Contents Preface xvii About the Authors DNA Function II: Protein Synthesis 39 DNA Structure II: Chromosomes and Cell Division 45 xxiii Part I Foundations 1 What Is Biological Anthropology? Anthropology and Its Subfields 1 INNOVATIONS: DNA Barcoding 56 3 The Scope of Biological Anthropology Summary • Review Questions 3 The Subfields of Anthropology 4 Paleoanthropology 5 Skeletal Biology and Human Osteology Paleopathology and Bioarchaeology 6 Forensic Anthropology 7 Primatology 8 Human Biology 9 4 6 10 Summary • Review Questions Origins of Evolutionary Thought What Is Science? 12 14 The Early Thinkers 15 The Roots of Modern Science 15 Linnaeus and the Natural Scheme of Life 16 The Darwinian Revolution 19 The Galápagos 19 Refining the Theory of Evolution by Natural Selection 22 Heritability and IQ Test Score Performance 26 Summary • Review Questions Genetics: Cells and Molecules INNOVATIONS: A New Genetic Era Genes and Environments 5 30 32 34 INSIGHTS & ADVANCES: Cloning Controversies DNA Structure and Function 37 DNA Structure I: The Molecular Level DNA Function I: Replication 39 27 79 80 Summary • Review Questions PART II Mechanisms of Evolution The Cell 33 Cell Anatomy 77 Phenylketonuria: Illustrating Mendelian and Post-Mendelian Concepts 78 24 INSIGHTS & ADVANCES: What Is Intelligent Design? The Study of Genetics 68 Mutation 68 Point Mutation and Sickle Cell Disease 68 Mutations: Bad, Neutral, and Good 70 X-Linked Disorders 72 Mendelian Genetics in Humans 73 INSIGHTS & ADVANCES: Popular Mendelism and the Shadow of Eugenics 76 INSIGHTS & ADVANCES: Darwin versus Wallace? 3 Mendelian Genetics 63 Mendel’s Postulates 65 Linkage and Crossing Over Genetics beyond Mendel 73 Polygenic Traits, the Phenotype, and the Environment 75 The Road to the Darwinian Revolution 16 The Uniformitarians: Hutton and Lyell 18 Science and Creationism Genetics: From Genotype to Phenotype 59 From Genotype to Phenotype 61 The ABO Blood Type System 62 Obesity: A Complex Interaction 62 The Roots of Modern Biological Anthropology 2 Molecular Tools for Bioanthropological Research 51 Indirect and Direct Research Methods 51 PCR, Mitochondrial DNA, and Ancient DNA 52 37 36 The Forces of Evolution and the Formation of Species 83 How Evolution Works 84 Where Does Variation Come From? 84 How Natural Selection Works 85 Other Ways By Which Evolution Happens 87 Classification and Evolution 91 Taxonomy and Speciation 91 What Is a Species? 92 Species Concepts 94 xi xii Contents INSIGHTS & ADVANCES: What’s in a Name? Species Concepts, Genetics, and Conservation 95 Life History Traits 148 Behavioral Traits 149 A Guide to the Nonhuman Primates The Strepsirhines 154 The Haplorhines 157 Reproductive Isolating Mechanisms 96 How Species Are Formed 96 The Tempo of Speciation 98 Adaptation 98 Is Everything Adaptive? 99 Hardy–Weinberg Equilibrium INSIGHTS & ADVANCES: The Rarest of the Rare 100 Primate Ecology Summary • Review Questions 105 Human Variation at the Individual and Group Levels What Is a Population? 107 107 Historical Perspectives on Human Variation 108 Recording Human Variation in Past Civilizations 108 The Monogenism–Polygenism Debate 110 Race and Racism in the Twentieth Century 111 Changing Attitudes toward Race in Anthropology 112 Deconstructing Racial Features 112 182 186 188 191 INSIGHTS & ADVANCES: The Infanticide Wars 194 INSIGHTS & ADVANCES: Are Chimpanzees from Mars and Bonobos from Venus? 196 Summary • Review Questions PART IV Body Size and Shape 132 Living at High Altitude 133 Skin Color 135 9 The Fossil Record Geology and Primate Origins 200 How to Become a Fossil Summary • Review Questions 202 The Importance of Context 203 Stratigraphy 203 The Geologic Time Scale 205 Primates 141 144 178 Types of Nonhuman Primate Societies INSIGHTS & ADVANCES: Technology and Extreme Environments 131 What Exactly Is a Primate? Anatomical Traits 144 Primate Behavior INNOVATIONS: Culture in Nonhuman Primates Adaptation and Adaptability 129 Levels of Adaptability 129 Heat and Cold 130 The Primate Radiation 143 The Extraordinary Diversity of Nonhuman Primates 8 Why Are Nonhuman Primates Social? The Paradox of Sociality 187 121 Polymorphisms and Natural Selection in Human Populations 123 The Evolution of Lactose Tolerance 123 Balanced Polymorphisms: Sickle Cell and Other Conditions 125 The Primates Summary • Review Questions The Evolution of Primate Social Behavior 181 Social Behavior and Reproductive Asymmetry Male Reproductive Strategies 184 Female Reproductive Strategies 185 INNOVATIONS: Bermuda: Population Genetics of One of the Last Places Settled by People 120 7 Diet 171 You Are What You Eat: Dietary and Digestive Strategies 172 Diet and Feeding Competition 173 Territories and Ranges 173 Predation 174 Primate Communities 175 Studying Primates 180 Population Genetics 114 Polymorphisms: ABO and Other Blood Type Systems 114 Gene Flow and Protein Polymorphisms 118 PART III 169 INSIGHTS & ADVANCES: The Impending Extinction of the Great Apes? 170 Human Variation: Evolution, Adaptation, and Adaptability Polymorphisms and Phylogenetic Studies 158 The New World Monkeys 159 The Old World Monkeys 161 The Hominoids 163 Levels of Selection 101 Inclusive Fitness 102 6 150 143 How Old Is It? 208 Relative Dating Techniques 209 Calibrated Relative Dating Techniques 211 Chronometric Dating Techniques 212 INSIGHTS & ADVANCES: Dating Controversies INNOVATIONS: Time in a Bottle 216 215 Contents 11 Earth in the Cenozoic 219 Continents and Land Masses 219 The Environment in the Cenozoic 220 Climate Change and Early Primate Evolution 222 Changes in the Paleocene Related to the Origin of Primates 223 Why Primates? 224 True Primates of the Eocene 224 Selective Pressures Favoring the Strepsirhine–Haplorhine Split 226 Climate Change and the Origin of Monkeys and Apes 227 The First Monkeys 227 New World Monkeys 229 Old World Monkeys 230 What Favored the Origin of Anthropoids? 230 The Earliest Apes 231 Selection Pressures and the Divergence of Monkeys and Apes 233 The Monkey’s Tale: What Happened to Primate Diversity in the Miocene? 236 277 Climate and the Evolution of Homo in the Pliocene and Pleistocene 278 Defining the Genus Homo 279 Earliest Genus Homo 279 Homo habilis (1.9–1.4 mya) 280 Homo rudolfensis (2.1–1.78 mya) 281 Early Tool Use, Hunting, and Scavenging 281 INSIGHTS & ADVANCES: Homo naledi and The Cradle of Humankind 282 Hunting and Scavenging 285 Who Was Homo erectus? 286 Anatomical Features 286 Homo erectus versus Homo ergaster 290 Homo erectus Around the World 291 African Origins 291 The First African Diaspora: Republic of Georgia Dispersal Into East Asia 295 INNOVATIONS: What’s Size Got to Do with It? Molecular Evolution in Primates 236 A Primate Molecular Phylogeny 237 The Status of Homo erectus in Europe The Lifeways of Homo erectus 301 Homo erectus and the Early Stone Age Summary • Review Questions 10 Rise of the Genus Homo 294 298 300 301 INSIGHTS & ADVANCES: The Little People of Flores Early Hominins and Australopithecus 242 Becoming a Biped 243 Anatomical Changes 244 Constructing the Bipedal Body Plan 302 A Higher-Quality Diet: Homo erectus Subsistence 304 Homo Erectus Life History 304 Homo erectus Leaves Africa 305 Summary • Review Questions 247 Will You Know a Hominin When You See One? 248 The First Hominins? 249 Sahelanthropus tchadensis (7.0–6.0 mya) 249 INSIGHTS & ADVANCES: Locomotion of the Last Common Ancestor 250 Orrorin tugenensis (6.0 mya) 252 Ardipithecus ramidus (4.4 mya) and Ardipithecus kadabba (5.8–5.2 mya) 253 Selective Pressures and the Origin of Hominins 254 Australopithecus and Kin 257 Australopithecus anamensis (4.2–3.9 mya) 257 Australopithecus afarensis (3.9–2.9 mya) 260 INNOVATIONS: Dikika and Development 262 Other East and West African Hominins (3.5–2.5 mya) 264 Australopithecus africanus (3.5–
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