Matt Cartmill is Professor of Anthropology at Boston University and Professor Emeritus of Evolutionary Anthropology at Duke University. Dr Cartmill is a Guggenheim and AAAS Fellow, a former president of the American Association of Physical Anthropologists and recipient of their Charles Darwin Lifetime Achievement Award, a founding co-editor of the International Journal of Primatology, and the former editor-in-chief of the American Journal of Physical Anthropology.Fred H. Smith is University Professor Emeritus of Anthropology at Illinois State University and Adjunct Professor at the University of Colorado, Boulder. A past president of the American Association of Physical Anthropologists and an AAAS and Alexander von Humboldt Fellow, he has received awards for his work from the U.S., Ireland, Germany and Croatia. Dr. Smith has conducted research in Europe, West Asia, and Africa, and has taught internationally at the Universities of Hamburg, Tübingen and Zagreb.

• Foreword xiPreface to the First Edition xiiiPreface to the Second Edition xviSome Notes on Nomenclature xixAbout the Companion Website xxi1 The Fossil Record 11.1 The Discovery of the Deep Past 1Changing Ideas About the Changing Earth 1Neptune vs. Vulcan 2A Brief Guide to Sedimentology 3Dating the Rocks 4The Succession of Faunas 5Radiation-Based Dating Techniques 7Other Dating Techniques 9Dating Based on the Cycles of the Earth 9The Problem of Orogeny 11Continental Drift 111.2 A Brief History of Life 12Life: The First Three Billion Years 12Multicellular Life 14The Cambrian Revolution 15Jaws, Fins, and Feet 16The Reptilian Revolutions 18The Two Great Extinctions 20The Mammals Take Over 212 Analyzing Evolution 232.1 Darwin and Evolution 23Parsimony and Pigeons 23Darwin’s Theory 24Improving on Darwin 272.2 The Origin of Species 30What, if Anything, is a Species? 30The Speciation Process 31The Tempo of Speciation 32Semispecies, Hybrids, and Isolating Mechanisms 332.3 Species Concepts and Classification 35Races, Semispecies, and Taxonomy 35Other Species Concepts 37Morphospecies and Chronospecies 392.4 Microevolution and Macroevolution 40Is Evolution Smooth or Jerky? 40The Neo-Darwinian Synthesis 41The Politics of Macroevolution 422.5 Reconstructing the Tree of Life 42Phylogenetic Inference 42Sources of Error in Phylogenetics 442.6 Taxonomy and Classification 47Linnaean Systematics 47Evolutionary Systematics 47Phenetics and Cladistics 49Pros and Cons of Phylogenetic Systematics 493 People as Primates 513.1 Primates as Mammals 51The First Mammals 51Allometry 57Allometry and Early Mammals 58Death and Molar Occlusion 59Allometry, Motherhood, and Milk 60Respiration and the Palate 60The Tribosphenic Molar 62Live Birth and Placentation 64Jurassic and Cretaceous Mammals 653.2 The Order Primates 66What is a Primate? 66The Living Strepsirrhines 73Anthropoid Apomorphies: Ears, Eyes, and Noses 74Tarsiers 76Platyrrhines: The New World Anthropoids 77Cercopithecoids: The Old World Monkeys 78Hominoids: The Living Apes 79Pongids and Hominids 81Bonobos and Chimpanzees 84Humans vs. Apes: Skulls and Teeth 853.3 The Primate Fossil Record 88Primate Origins: The Crown Group 88Fossil Primates: The Stem Group 90Ancestral Traits and Genetic Evidence 91The First Euprimates 92Eocene “Lemurs” and “Tarsiers” 94The First Anthropoids 96Anthropoid Radiations 98Miocene Catarrhines 99Ape Origins 103Cercopithecoids 1074 The Bipedal Ape 1094.1 The Discovery of Australopithecus 109Being Human vs. Becoming Human 109The Taung Child 109Australopithecus Grows Up 1114.2 The Anatomy of Bipedality 115Upright Posture and the Vertebral Column 115Bipedality and the Pelvis 116Bipedal Locomotion: Knees 118Bipedal Locomotion: The Hip Joint 123Bipedal Locomotion: Feet 1244.3 More South African Finds 127Australopithecus Stands Up 127The Skull of Australopithecus africanus 128Australopithecus robustus 129Man-Apes, Just Plain Apes, or Weird Apes? 133Postcranial Peculiarities 1334.4 Louis Leakey and Olduvai Gorge 1354.5 Mio-Pliocene Enigmas 139Sahelanthropus: The Oldest Hominin? 139Orrorin 140Ardipithecus 141The Burtele Foot 1464.6 The Genus Australopithecus 146Australopithecus anamensis? 146Australopithecus afarensis? 148Afarensis Skulls and Teeth 152Australopithecus bahrelghazali? 153Australopithecus deyiremeda? 153Kenyanthropus platyops? 154Early Australopithecus from South Africa 154Australopithecus prometheus? 155Australopithecus aethiopicus 156Australopithecus garhi 158Australopithecus sediba? 159Australopithecus boisei 160Australopithecus robustus: Postcranial Skeleton and Relationships 1624.7 Australopithecine Phylogeny 163Alpha Taxonomy and Cladograms 163Getting Around Cladistics 1664.8 The Australopithecine Postcranium 167Down from the Trees – How Far, How Fast? 167Australopithecine Shoulders 171Arms vs. Legs 172The Hominin Hand 173Australopithecine Vertebrae 174Hip and Femur 177Early Hominin Feet 179Postcranial Diversity in Early Hominins 1814.9 Ecology and Behavior 183The Facts Thus Far 183What Did Australopithecines Eat? 183Early Hominin Environments 186Social Ecology 1884.10 Major Issues: Explaining Hominin Origins 1925 The Migrating Ape 1975.1 The Spread of Hominins out of Africa 1975.2 The Emergence of the Genus Homo 198Homo habilis and the Habilines 198Habiline Dates and Stratigraphy in East Africa 203Habiline Skulls 204Habiline Teeth and Diets 207Habiline Postcranial Remains 208Habiline Taxonomy: The Frustrations of Variation 210Back to South Africa 211Advanced Australopithecus or Early Homo? Phylogenetic Issues 212Early Material Culture 214A Summary of the Habilines in Eight Questions 2155.3 Homo erectus 216An Introduction to Homo erectus 216A Brief History of Homo erectus: 1889–1950 218Later Discoveries in Africa and Eurasia 220Erectine Chronology and Geographic Distribution 222Asian Homo erectus: The Neurocranium 224Cranial Capacity and the Brain in Asian Erectines 229Asian Homo erectus: Faces and Mandibles 230The Asian Erectine Dentition 232Asian Erectine Postcranial Remains 233Early African Erectine Skulls and the Ergaster Question 233Early African Erectine Postcranial Morphology 237Early Erectine Adaptations: Anatomy and Physiology 242Early Erectine Adaptations: The Archaeological Evidence 245Patterns of Development and Evolutionary Change in Erectines 247Early Erectine Radiations in Africa 248Out of Africa I: The First Migration into Eurasia 250Dmanisi – The First Eurasians 253Indonesian Erectines and the Specter of “Meganthropus” 259Chinese Erectines 261The Initial Occupation of Europe 262Gran Dolina 2645.4 Peripheral Holdouts along the Continental Margins 267Flores 267Luzon 272Dushan 272Rising Star 2735.5 Major Issues: Summing Up the Erectines 2756 The Big-Brained Ape: Middle Pleistocene Variants and Trends 2796.1 Homo “heidelbergensis” 279Crossing the Rubicon? 279“Archaic Homo sapiens” vs. “Homo heidelbergensis” 280Brains and Tools in the Middle Pleistocene 2826.2 Models of Later Human Evolution 284Changing Origin Narratives 284The Piltdown Fraud 285RAO and MRE 2876.3 Regional Variants in Europe and Africa 289European Heidelbergs 289Petralona 290Bilzingsleben 293Swanscombe 293Steinheim 294Mauer 295Boxgrove 296Ceprano 296Arago (Tautavel) and Lazaret 297Sima de los Huesos 298Other European Heidelbergs 303African Heidelbergs: Kabwe 304Bodo and Ndutu 306African Heidelberg Mandibles 307Other African Heidelbergs 307North Africans 3086.4 Asian Heidelbergs? 308Mugharet El-Zuttiyeh 308Other West Asian Candidates 309South Asia 309East Asia 3096.5 Australasia 311Sambungmacan 311Ngandong 3126.6 Supraorbital Tori, Chins, and Projecting Faces 3146.7 The African Transition to Modern Humans 316Background and Dating 316The African Transitional Group: Vault Morphology 319The African Transitional Group: Facial Morphology 320The African Transitional Group: Additional Bones, Archaeology, and Other Matters 3216.8 East Asian Archaic Humans 322Background and Context 322Dali 324Harbin 325Other Chinese Finds 325East Asian Archaics: Continuity or Someone New? 3266.9 Major Issues: Speciation, Migration, and Regional Differentiation 3277 Talking Apes: The Neandertals 3337.1 Changing Ideas about Neandertals 333Early Discoveries and Interpretations 335Neandertals – From Boule to the Twenty-First Century 3387.2 Neandertal Chronology and Distribution 3407.3 The Neandertal Skull 346Neandertal Braincases 346Neandertal Faces 356Neandertal Mandibles 359Neandertal Teeth 362Prognathism 3647.4 The Neandertal Body 366Body Size and Proportions 366Neck and Upper Limb 369Lower Vertebrae, Pelvis, and Lower Limb 3717.5 Neandertal Life History and Demography 3747.6 Genetics and Genomics 376The Mitochondrial Genome 376The Nuclear Genome 378Genes, Dates, and Lineages 379Denisovans 380Ghosts in the Genes 3827.7 Brains and Behavior 383Neandertal Brains 383Neandertal Technology 385Symbolic Behavior 386Neandertals and Language 389Diets and Subsistence Behavior 3937.8 Neandertal Populations 395Early European Neandertals 395Krapina 396“Würm” Neandertals from Western Europe 398Western and Central Asian Neandertals 399Late Neandertals 4027.9 Major Issues 4068 The Symbolic Ape: The Origins of Modern Humans 4118.1 Symbolic Behavior 411Signs and Symbols 411A “Creative Explosion”? 4128.2 Modern Human Anatomy 414The Modern Skull 414Cranial Capacity 416The Postcranial Skeleton 4178.3 The Fossil Record of Modern Human Origins 418Geochronology 418Early Modern Humans: The East African Record 418Out of (East) Africa: Early Modern People in North and South Africa 421The First Modern People Outside Africa: The Near Eastern Evidence 424African and Circum-Mediterranean Gene Flow and Modern Human Origins 430Modern Human Origins in East Asia 432The First Australians 437Europe: A Late Frontier 443The Initial Upper Paleolithic 444The Aurignacian and its Makers 445The Gravettian 450The Late Entry into Europe 452Europe: The Morphological Evidence for Continuity 4538.4 Genetics and Modern Human Origins 455Genes, Populations, and Migrations 455Human Self-Domestication? 459Ancient DNA in Early Modern Humans 4608.5 Modern Human Origins: The Models vs. the Data 461The Recent African Origin Model 461Multiregional Evolution 462Alternative Views: The Assimilation Model 463Assimilation and Interactions Between Modern and Archaic Humans 466Appendix: Cranial Measurements 471Bibliography 477Index 583