The Human Lineage, Second Edition: Foundation of Human Biology

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 xi Preface to the First Edition xiii Preface to the Second Edition xvi Some Notes on Nomenclature xix About the Companion Website xxi 1 The Fossil Record 1 1.1 The Discovery of the Deep Past 1 Changing Ideas About the Changing Earth 1 Neptune vs. Vulcan 2 A Brief Guide to Sedimentology 3 Dating the Rocks 4 The Succession of Faunas 5 Radiation-Based Dating Techniques 7 Other Dating Techniques 9 Dating Based on the Cycles of the Earth 9 The Problem of Orogeny 11 Continental Drift 11 1.2 A Brief History of Life 12 Life: The First Three Billion Years 12 Multicellular Life 14 The Cambrian Revolution 15 Jaws, Fins, and Feet 16 The Reptilian Revolutions 18 The Two Great Extinctions 20 The Mammals Take Over 21 2 Analyzing Evolution 23 2.1 Darwin and Evolution 23 Parsimony and Pigeons 23 Darwin's Theory 24 Improving on Darwin 27 2.2 The Origin of Species 30 What, if Anything, is a Species? 30 The Speciation Process 31 The Tempo of Speciation 32 Semispecies, Hybrids, and Isolating Mechanisms 33 2.3 Species Concepts and Classification 35 Races, Semispecies, and Taxonomy 35 Other Species Concepts 37 Morphospecies and Chronospecies 39 2.4 Microevolution and Macroevolution 40 Is Evolution Smooth or Jerky? 40 The Neo-Darwinian Synthesis 41 The Politics of Macroevolution 42 2.5 Reconstructing the Tree of Life 42 Phylogenetic Inference 42 Sources of Error in Phylogenetics 44 2.6 Taxonomy and Classification 47 Linnaean Systematics 47 Evolutionary Systematics 47 Phenetics and Cladistics 49 Pros and Cons of Phylogenetic Systematics 49 3 People as Primates 51 3.1 Primates as Mammals 51 The First Mammals 51 Allometry 57 Allometry and Early Mammals 58 Death and Molar Occlusion 59 Allometry, Motherhood, and Milk 60 Respiration and the Palate 60 The Tribosphenic Molar 62 Live Birth and Placentation 64 Jurassic and Cretaceous Mammals 65 3.2 The Order Primates 66 What is a Primate? 66 The Living Strepsirrhines 73 Anthropoid Apomorphies: Ears, Eyes, and Noses 74 Tarsiers 76 Platyrrhines: The New World Anthropoids 77 Cercopithecoids: The Old World Monkeys 78 Hominoids: The Living Apes 79 Pongids and Hominids 81 Bonobos and Chimpanzees 84 Humans vs. Apes: Skulls and Teeth 85 3.3 The Primate Fossil Record 88 Primate Origins: The Crown Group 88 Fossil Primates: The Stem Group 90 Ancestral Traits and Genetic Evidence 91 The First Euprimates 92 Eocene "Lemurs" and "Tarsiers" 94 The First Anthropoids 96 Anthropoid Radiations 98 Miocene Catarrhines 99 Ape Origins 103 Cercopithecoids 107 4 The Bipedal Ape 109 4.1 The Discovery of Australopithecus 109 Being Human vs. Becoming Human 109 The Taung Child 109 Australopithecus Grows Up 111 4.2 The Anatomy of Bipedality 115 Upright Posture and the Vertebral Column 115 Bipedality and the Pelvis 116 Bipedal Locomotion: Knees 118 Bipedal Locomotion: The Hip Joint 123 Bipedal Locomotion: Feet 124 4.3 More South African Finds 127 Australopithecus Stands Up 127 The Skull of Australopithecus africanus 128 Australopithecus robustus 129 Man-Apes, Just Plain Apes, or Weird Apes? 133 Postcranial Peculiarities 133 4.4 Louis Leakey and Olduvai Gorge 135 4.5 Mio-Pliocene Enigmas 139 Sahelanthropus: The Oldest Hominin? 139 Orrorin 140 Ardipithecus 141 The Burtele Foot 146 4.6 The Genus Australopithecus 146 Australopithecus anamensis? 146 Australopithecus afarensis? 148 Afarensis Skulls and Teeth 152 Australopithecus bahrelghazali? 153 Australopithecus deyiremeda? 153 Kenyanthropus platyops? 154 Early Australopithecus from South Africa 154 Australopithecus prometheus? 155 Australopithecus aethiopicus 156 Australopithecus garhi 158 Australopithecus sediba? 159 Australopithecus boisei 160 Australopithecus robustus: Postcranial Skeleton and Relationships 162 4.7 Australopithecine Phylogeny 163 Alpha Taxonomy and Cladograms 163 Getting Around Cladistics 166 4.8 The Australopithecine Postcranium 167 Down from the Trees - How Far, How Fast? 167 Australopithecine Shoulders 171 Arms vs. Legs 172 The Hominin Hand 173 Australopithecine Vertebrae 174 Hip and Femur 177 Early Hominin Feet 179 Postcranial Diversity in Early Hominins 181 4.9 Ecology and Behavior 183 The Facts Thus Far 183 What Did Australopithecines Eat? 183 Early Hominin Environments 186 Social Ecology 188 4.10 Major Issues: Explaining Hominin Origins 192 5 The Migrating Ape 197 5.1 The Spread of Hominins out of Africa 197 5.2 The Emergence of the Genus Homo 198 Homo habilis and the Habilines 198 Habiline Dates and Stratigraphy in East Africa 203 Habiline Skulls 204 Habiline Teeth and Diets 207 Habiline Postcranial Remains 208 Habiline Taxonomy: The Frustrations of Variation 210 Back to South Africa 211 Advanced Australopithecus or Early Homo? Phylogenetic Issues 212 Early Material Culture 214 A Summary of the Habilines in Eight Questions 215 5.3 Homo erectus 216 An Introduction to Homo erectus 216 A Brief History of Homo erectus: 1889-1950 218 Later Discoveries in Africa and Eurasia 220 Erectine Chronology and Geographic Distribution 222 Asian Homo erectus: The Neurocranium 224 Cranial Capacity and the Brain in Asian Erectines 229 Asian Homo erectus: Faces and Mandibles 230 The Asian Erectine Dentition 232 Asian Erectine Postcranial Remains 233 Early African Erectine Skulls and the Ergaster Question 233 Early African Erectine Postcranial Morphology 237 Early Erectine Adaptations: Anatomy and Physiology 242 Early Erectine Adaptations: The Archaeological Evidence 245 Patterns of Development and Evolutionary Change in Erectines 247 Early Erectine Radiations in Africa 248 Out of Africa I: The First Migration into Eurasia 250 Dmanisi - The First Eurasians 253 Indonesian Erectines and the Specter of "Meganthropus" 259 Chinese Erectines 261 The Initial Occupation of Europe 262 Gran Dolina 264 5.4 Peripheral Holdouts along the Continental Margins 267 Flores 267 Luzon 272 Dushan 272 Rising Star 273 5.5 Major Issues: Summing Up the Erectines 275 6 The Big-Brained Ape: Middle Pleistocene Variants and Trends 279 6.1 Homo "heidelbergensis" 279 Crossing the Rubicon? 279 "Archaic Homo sapiens" vs. "Homo heidelbergensis" 280 Brains and Tools in the Middle Pleistocene 282 6.2 Models of Later Human Evolution 284 Changing Origin Narratives 284 The Piltdown Fraud 285 RAO and MRE 287 6.3 Regional Variants in Europe and Africa 289 European Heidelbergs 289 Petralona 290 Bilzingsleben 293 Swanscombe 293 Steinheim 294 Mauer 295 Boxgrove 296 Ceprano 296 Arago (Tautavel) and Lazaret 297 Sima de los Huesos 298 Other European Heidelbergs 303 African Heidelbergs: Kabwe 304 Bodo and Ndutu 306 African Heidelberg Mandibles 307 Other African Heidelbergs 307 North Africans 308 6.4 Asian Heidelbergs? 308 Mugharet El-Zuttiyeh 308 Other West Asian Candidates 309 South Asia 309 East Asia 309 6.5 Australasia 311 Sambungmacan 311 Ngandong 312 6.6 Supraorbital Tori, Chins, and Projecting Faces 314 6.7 The African Transition to Modern Humans 316 Background and Dating 316 The African Transitional Group: Vault Morphology 319 The African Transitional Group: Facial Morphology 320 The African Transitional Group: Additional Bones, Archaeology, and Other Matters 321 6.8 East Asian Archaic Humans 322 Background and Context 322 Dali 324 Harbin 325 Other Chinese Finds 325 East Asian Archaics: Continuity or Someone New? 326 6.9 Major Issues: Speciation, Migration, and Regional Differentiation 327 7 Talking Apes: The Neandertals 333 7.1 Changing Ideas about Neandertals 333 Early Discoveries and Interpretations 335 Neandertals - From Boule to the Twenty-First Century 338 7.2 Neandertal Chronology and Distribution 340 7.3 The Neandertal Skull 346 Neandertal Braincases 346 Neandertal Faces 356 Neandertal Mandibles 359 Neandertal Teeth 362 Prognathism 364 7.4 The Neandertal Body 366 Body Size and Proportions 366 Neck and Upper Limb 369 Lower Vertebrae, Pelvis, and Lower Limb 371 7.5 Neandertal Life History and Demography 374 7.6 Genetics and Genomics 376 The Mitochondrial Genome 376 The Nuclear Genome 378 Genes, Dates, and Lineages 379 Denisovans 380 Ghosts in the Genes 382 7.7 Brains and Behavior 383 Neandertal Brains 383 Neandertal Technology 385 Symbolic Behavior 386 Neandertals and Language 389 Diets and Subsistence Behavior 393 7.8 Neandertal Populations 395 Early European Neandertals 395 Krapina 396 "Würm" Neandertals from Western Europe 398 Western and Central Asian Neandertals 399 Late Neandertals 402 7.9 Major Issues 406 8 The Symbolic Ape: The Origins of Modern Humans 411 8.1 Symbolic Behavior 411 Signs and Symbols 411 A "Creative Explosion"? 412 8.2 Modern Human Anatomy 414 The Modern Skull 414 Cranial Capacity 416 The Postcranial Skeleton 417 8.3 The Fossil Record of Modern Human Origins 418 Geochronology 418 Early Modern Humans: The East African Record 418 Out of (East) Africa: Early Modern People in North and South Africa 421 The First Modern People Outside Africa: The Near Eastern Evidence 424 African and Circum-Mediterranean Gene Flow and Modern Human Origins 430 Modern Human Origins in East Asia 432 The First Australians 437 Europe: A Late Frontier 443 The Initial Upper Paleolithic 444 The Aurignacian and its Makers 445 The Gravettian 450 The Late Entry into Europe 452 Europe: The Morphological Evidence for Continuity 453 8.4 Genetics and Modern Human Origins 455 Genes, Populations, and Migrations 455 Human Self-Domestication? 459 Ancient DNA in Early Modern Humans 460 8.5 Modern Human Origins: The Models vs. the Data 461 The Recent African Origin Model 461 Multiregional Evolution 462 Alternative Views: The Assimilation Model 463 Assimilation and Interactions Between Modern and Archaic Humans 466 Appendix: Cranial Measurements 471 Bibliography 477 Index 583