Advances in Human Biology t Matt Cartmill and Kaye Brown, Series Editors THIN ON THE GROUND Neandertal Biology, Archeology, and Ecology Steven E. Churchill JWST461-c01 JWST461-Churchill Printer: Yet to Come June 11, 2014 6:52 Trim: 254mm×178mm JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm Thin on the Ground JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm Advances in Human Biology Series Editors: Matt Cartmill Kaye Brown Boston University Titles in this Series Thin on the Ground: Neandertal Biology, Archeology, and Ecology by Steven E. Churchill JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm THIN ON THE GROUND Neandertal Biology, Archeology, and Ecology STEVEN EMILIO CHURCHILL Department of Evolutionary Anthropology Duke University USA SERIES EDITORS: MATT CARTMILL AND KAYE BROWN JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm This edition first published 2014 © 2014 by John Wiley & Sons, Inc. Editorial offices: 1606 Golden Aspen Drive, Suites 103 and 104, Ames, Iowa 50010, USA The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 9600 Garsington Road, Oxford, OX4 2DQ, UK For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. 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Cover drawing by Matt Cartmill Set in 10/12pt Garamond by Aptara Inc., New Delhi, India 1 2014 JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm Contents 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 v JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm vi Contents 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 JWST461-fm JWST461-Churchill Printer: Yet to Come July 7, 2014 7:14 Trim: 254mm×178mm Contents vii 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 ....................