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1 1 low Humans Evolved Robert Howl IOIIIB Silk UNIVERS1. i 1 \..UK I..1 I \ Nv I Technische Unive-^itdt Darmstadt FACHDCRLICH 10 BIOLOGIE B i!. I i o t h p k -_ ScLninspilinstiafiG 10 D Darmstadt inv.-nr. \\ \\ Nmtoii «.\ ( ompiny \ oik London

2 Conlcnls 1'iolo.u \,\ i\ SUMl\ llunii'i I liiimn* Part One: How Evolution Works Chapter 1: AdaptatiorubyjNatinal Selection Explaining- Adaptation beton. Dai win I Darwin's ^Theory of Adaptation 4 N* V ^ Darwiii'-s Postulates S fffil^y- An Example v 'of Adaptation In Natural Seleition,";;4.'"4 Individual Selection 14 "/T-^/Fhe, Evolution of Complex Ad.ipt.itions 16 <1 "}-'Why Small Variations -\ic Important U> «-; Why IntermediijeJ^leps Aie f-.ivored In S(.leition Rates of Eyolutionary^iiaiii;! )aiv\ ID'S I )itfkulties Expl.imuiii Vatiation!( > I IS Chapter 2: Cienetus 29 Mcndfli.in ;^ Cell Division and the"role ol Chromosomes m Inheiiunc i- Mitosis and-,meiosis V3 Chromosomes and'mendel's r\pennionral Results 35 Linkage and Recombiiution 40 Molecular Genetics 4f>.. Genes Are DNA 4~ The Chemical Basis., of Life ^<' DMA Codes for Protein ^2 Not All DNA C lodes loi Proteins SS

3 Chapter 3: The Modern Synthesis 63 Population Genetics 63 Genes in Populations 64 How Random Mating and Sexual Reproduction Change Genotypic Frequencies 65 How Natural Selection Changes Gene Frequencies 70 The Modern Synthesis 72 The Genetics of Continuous Variation 72 How Variation Is Maintained 78 Natural Selection and Behavior 82 Constraints on Adaptation 86 Correlated Characters 86 Disequilibrium 89 Genetic Drift 90 Local versus Optimal Adaptations 95 Other Constraints on Evolution 97 Chapter 4: Speciation and Phylogeny 103 What Are Species? 103 The Biological Species Concept 105 The Ecological Species Concept 107 The Origin of Species 109 Allopatric Speciation 110 Parapatric and Sympatric Speciation 113 The Tree of Life 121 Why Reconstruct Phylogenies? 124 How to Reconstruct Phylogenies 130 Problems Due to Convergence 131 Problems Due to Ancestral Characters 132 Reconstructing Phylogenies Using Genetic Distance Data 136 Taxonomy Naming Names 141 Part Two: Primate Behavior and Ecology Chapter 5: Introduction to the Primates 147 Two Reasons to Study the Primates 147 Primates Are Our Closest Relatives 148 Primates Are a Diverse Order 148 Features That Define the Primates 149 Primate Biogeography 156 A Taxonomy of Living Primates 157 The Prosimians 158 The Anthropoids 164 Primate Conservation 174

4 Chapter 6: Primate Ecology 179 The Distribution of Food 180 Activity Patterns 193 Ranging Behavior 195 Predation 199 Chapter 7: Primate Mating Systems 203 The Language of Adaptive Explanations 205 The Evolution of Reproductive Strategies 206 Reproductive Strategies of Primate Females 209 Female Dominance Hierarchies 212 Reproductive Tradeoffs 215 Primate Sociality 219 Why Do Primates Live in Groups? 220 How Big Should Groups Be? 225 What Kinds of Groups Should Primates Form? 226 Sexual Selection and Male Mating Strategies 231 Intrasexual Selection in Primates 234 Intersexual Selection in Primates 236 Sexual Selection and Primate Behavior 239 Monogamous Males 239 One-Male, Multifemale Groups 241 Multimale, Multifemale Groups 247 Chapter 8: The Evolution of Social Behavior 253 Kinds of Social Interactions 254 Altruism: A Conundrum 255 Kin Selection 258 Hamilton's Rule 260 Evidence of Kin Selection in Primates 263 Reciprocal Altruism 271 Chapter 9: Primate Intelligence 281 What Is Intelligence? 281 Why Are Primates So Smart? 283 Hypotheses Explaining Primate Intelligence 284 Testing Models of the Evolution of Intelligence 286 The Value of Studying Primate Behavior 307 Part Three: The History of the Human Lineage Chapter 10: From Tree Shrew to Ape 311 Continental Drift and Climate Change 314 The Methods of Paleontology 317

5 The Evolution of the Early Primates 319 The First Anthropoids 324 The Emergence of the Hominoids 334 Chapter 11: The Earliest Hominids 342 Australopithecus afarensis 343 Morphology 346 Bipedal Locomotion 347 Other Features of A. afarensis 356 When Is a Hominoid Also a Hominid? 357 Hominids before A. afarensis 359 Australopithecines after A. afarensis 362 A. africanus 362 Robust Australopithecines 365 Early Homo 373 Traits Defining Early Homo 374 How Many Species? 375 Flaked Stone Tools 377 Hominid Phylogenies 386 Before Discovery of WT After Discovery of WT Chapter 12: The Lives of Early Hominids 392 From Ape to Human 393 Early Hominid Environments 396 Early Hominid Ecology The Emergence of Meat Eating 397 Why Meat Eating Is Important 398 Comparison with Hunting in Chimpanzees 399 Seasonality and Meat Eating 401 Archaeological Evidence in Early Hominids 403 Hunters or Scavengers? 412 Early Hominid Social Organization 416 Sexual Division of Labor 417 Food Sharing 420 Archaeological Evidence 421 Chapter 13: The Ancients 427 Homo erectus 428 Acheulean Tools 435 Homo erectus Peoples the World 437 j Cultural Adaptations of H. erectus 440 Archaic Homo sapiens 448 The Neanderthals and Their Contemporaries 450 Mousterian and Middle Stone Age Tools 456 Neanderthal Lifeways 458

6 Chapter 14: The Moderns 464 The Emergence of Anatomically Modern People 464 The Upper Paleolithic the Human Revolution 467 Technology and Culture 471 Symbolic Behavior 474 Subsistence and Social Organization 481 Models for the Origin and Spread of Anatomically Modern Humans 486 Evidence from Fossils and Tool Kits 488 Genetic Data 491 Chapter 15: The Evolution of Language 500 Language Is an Adaptation 501 Speech Production and Perception 503 Grammar 506 Language Capacities Are Derived 512 How Language Evolved 515 When Language Evolved 519 Did Language Arise Early? 519 Did Language Arise Late? 523 Part Four: Evolution and Modern Humans Chapter 16: Human Genetic Diversity 531 Explaining Human Variation 531 Variation in Traits Influenced by Single Genes 535 Causes of Genetic Variation within Groups 537 Causes of Genetic Variation among Groups 541 Variation in Complex Phenotypic Traits 549 Genetic Variation within Groups 550 Genetic Variation among Groups 552 The Race Concept 556 Chapter 17: Evolution and the Human Life Cycle 571 Maternal-Fetal Conflict during Pregnancy 572 Why There Is Parent-Offspring Conflict ; 572 Spontaneous Abortion 574 Blood Sugar 576 Placental Blood Supply 577 The Evolution of Senescence 580 Two Evolutionary Theories of Senescence 582 Evidence for the Theories 586 The Evolution of Menopause 588

7 Chapter 18: Evolution and Human Behavior 597 Why Evolution Is Relevant to Human Behavior 597 Evolutionary Psychology 600 The Logic of Evolutionary Psychology 600 Reasoning about Reciprocity 603 Evolutionary Psychology and Human Universals 607 Color Terms 607 Inbreeding Avoidance 613 Evolution and Human Culture 620 Culture Is a Derived Trait in Humans 624 Culture Is an Adaptation 630 Human Behavioral Ecology 634 Chapter 19: Human Mate Choice and Parenting 641 The Psychology of Human Mate Preferences 642 Some Social Consequences of Mate Preferences 648 Kipsigis Bridewealth 648 Nyinba Polyandry 650 Raising Children 653 Child Abuse and Infanticide 654 Cross-Cultural Patterns of Infanticide 655 Child Abuse in the United States and Canada 657 Adoption 658 Adoption in Oceania 659 Adoption in Industrialized Societies 661 Family Size 662 Is Human Evolution Over? 667 Epilogue: There Is Grandeur in this View of Life Appendix The Skeletal Anatomy of Primates Al Glossary A3 Credits A19 Index A25

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