Thin on the Ground Neandertal Biology, Archeology, and Ecology Foundation of Human Biology Series

Thin on the Ground: Neandertal Biology, Archeology and Ecology synthesizes the current knowledge about our sister species the Neandertals, combining data from a variety of disciplines to reach a cohesive theory behind Neandertal low population densities and relatively low rate of technological innovation. The book highlights and contrasts the differences between Neandertals and early modern humans and explores the morphological, physiological, and behavioral adaptive solutions which led to the extinction of the Neandertals and the population expansion of modern humans.

Written by a world recognized expert in physical anthropology, Thin on the Ground: Neandertal Biology, Archaeology and Ecology will be a must have title for anyone interested in the rise and fall of the Neandertals.

Series Introduction ix

Preface xi

Acknowledgements xv

Chapter 1 Thin on the Ground: Population Density and Technological Innovation 1

Note 7

Chapter 2 The Neandertals in Time and Space 9

2.1 Geographic and Temporal Boundaries 10

2.2 Defining the Neandertals 13

2.3 Neandertal DNA 27

2.4 Neandertal Taxonomy 30

2.5 Regional and Temporal Variation in Neandertal Morphology 30

2.6 The Evolutionary History of the Neandertals 32

Notes 39

Chapter 3 Neandertal Material Culture 41

3.1 Neandertal-associated Lithic Industries 42

3.2 Variation in the Eurasian Middle Paleolithic: Technology as Adaptive Interface 53

3.3 Composite Technology, and the Archeologically Less-visible Component of Technology 59

3.4 Subsistence Technology 61

3.5 Domestic Technology 67

Notes 69

Chapter 4 The Body Neandertal 71

4.1 Neandertal Body Size: Short but Massive 72

4.2 Body Composition: “Scaled Up” Inuit? 79

4.3 The Cost of Size: Feeding a Large Body and Large Brain 82

4.3.1 Nutrition and Somatic Maintenance: Neandertal Bodies were Energetically Costly 83

4.3.2 Nutrition and Reproduction: Were Neandertal Mothers Like Polar Bears? 92

4.3.3 Dietary Shortfalls: Hard and Lean Winters 97

4.4 The Benefits of Size: Neandertal Body Size in Ecological Context 99

Notes 104

Chapter 5 Surviving the Cold 107

5.1 How Cold Was It? 108

5.1.1 Cold–Temperate Conditions (MIS 5d-a and 3) 109

5.1.2 Cold Glacial Conditions (MIS 6 and 4) 111

5.2 Human Adaptation to the Cold 114

5.3 Cold Adaptation and Neandertal Morphology 117

5.3.1 The Survival Value of Neandertal Body Form 124

5.3.2 Craniofacial Morphology and Cold Adaptation 129

5.4 Physiological Solutions to Cold Stress 137

5.5 Cold Stress and Neandertal Behavior 141

5.5.1 Activity 141

5.5.2 Extrasomatic Heat Production and Conservation: Fire, Shelter and Clothing 142

5.6 Thermogenic Capacity and Cold Tolerance 146

5.7 The Neandertals Were Cold-adapted 148

Notes 149

Chapter 6 The Caloric Economy of Pleistocene Europe 151

6.1 Issues in the Reconstruction of Past Environments 152

6.2 Pleistocene Biomes of Europe and Western Asia 157

6.2.1 Interglacial Europe: Marine Isotope Stages 7 and 5e 159

6.2.2 Productivity and Edible Resources of Interglacial Environments 162

6.2.3 Temperate Interglacial Europe: Marine Isotope Stages 5d–a and 3 169

6.2.4 Cold Steppic Europe: Marine Isotope Stages 6 and 4 173

Notes 175

Chapter 7 Neandertals as Consumers 179

7.1 Analysis of Food Residues: The Macromammal Component of Neandertal Diet 181

7.2 Analysis of Food Residues: The Small Animal Data 187

7.3 Analysis of Food Residues: Macrobotanical Remains 191

7.4 Dental Wear and Food Residues on Teeth 194

7.5 Stable Isotope and Trace Element Analyses 200

7.6 The Thorny Issue of Cannibalism 214

7.7 The Trophic Ecology of Neandertals 216

Notes 218

Chapter 8 Red in Tooth and Claw: Neandertals as Predators 219

8.1 Neandertal Morphology and Predation 220

8.1.1 Scapular Glenoid Fossa 222

8.1.2 Elbow Joint Morphology 224

8.1.3 Superior Pubic Ramus Length 226

8.1.4 Humeral Diaphyseal Cross-sectional Geometry 226

8.1.5 Entheseal (Muscle Marking) Morphology 233

8.2 Neandertals as Close-range Predators 236

8.3 Prey Size, Hunting “Pack” Size, and Risk of Injury to Neandertal Hunters 245

8.4 Neandertal Hunting in Ecological Context 246

Notes 249

Chapter 9 In the Company of Killers: Neandertals as Carnivores 251

9.1 Large-bodied Carnivores of the Eurasian Late Pleistocene 253

9.2 The Members of the Eurasian Pleistocene Large-bodied Carnivore Guild 256

9.2.1 Homotherium latidens 256

9.2.2 Panthera leo spelaea 258

9.2.3 Panthera pardus 259

9.2.4 Crocuta crocuta spelaea 260

9.2.5 Hyaena hyaena 262

9.2.6 Canis lupus 262

9.2.7 Cuon alpinus 263

9.3 Competition within the Carnivore Guild 264

9.3.1 Exploitation Competition 265

9.3.2 Interference Competition 268

9.4 Neandertals Were Not the Socially-Dominant Members of the Carnivore Guild 271

9.5 Neandertal Ecology in the Context of Competition within the Carnivore Guild 276

Notes 284

Chapter 10 The Cost of Living in Ice Age Europe 287

10.1 Subsistence Organization and Mobility 289

10.2 Home Range Size 299

10.2.1 Lithic Raw Material Movement 300

10.2.2 Carnivore Models for Estimating Neandertal Mobility 302

10.3 Paleontological Reflections of Neandertal Mobility 309

10.4 The Energetic Cost of Mobility 316

10.5 The Energetic Cost of Domestic Activities 321

10.6 Neandertal Physical Activity Levels 324

Notes 331

Chapter 11 Neandertal Social Life, Life History, and Demography 333

11.1 Subsistence Labor Demands, Group Size, and Social Structure 336

11.2 Neandertal Life History 342

11.3 Neandertal Demography 347

Notes 352

Chapter 12 From Thin to Thick: The African MSA 353

12.1 Tipping the Scales on Population Growth 355

12.2 Culture Change in the Late MSA and Mousterian 360

Note 363

References 365

Index 445

Steven E. Churchill is Professor of Evolutionary Anthropology at Duke University. A recognized leader in the field of physical anthropology, Dr. Churchill is the author of more than 50 research articles on the morphology and behavior of Neandertals and early modern humans.