A Morphometric Investigation into the Origin(s) of Anatomically Modern Humans 9781841713441, 9781407325828

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A Morphometric Investigation into the Origin( s) of Anatomically Modem Humans

Phillip J. Habgood

BAR International Series 1176 2003

Published in 2019 by BAR Publishing, Oxford BAR International Series 1176 A Morphometric Investigation into the Origin(s) of Anatomically Modern Humans © Phillip J. Habgood and the Publisher 2003 The author’s moral rights under the 1988 UK Copyright, Designs and Patents Act are hereby expressly asserted. All rights reserved. No part of this work may be copied, reproduced, stored, sold, distributed, scanned, saved in any form of digital format or transmitted in any form digitally, without the written permission of the Publisher. ISBN 9781841713441 paperback ISBN 9781407325828 e-book DOI https://doi.org/10.30861/9781841713441 A catalogue record for this book is available from the British Library This book is available at www.barpublishing.com BAR Publishing is the trading name of British Archaeological Reports (Oxford) Ltd. British Archaeological Reports was first incorporated in 1974 to publish the BAR Series, International and British. In 1992 Hadrian Books Ltd became part of the BAR group. This volume was originally published by John and Erica Hedges in conjunction with British Archaeological Reports (Oxford) Ltd / Hadrian Books Ltd, the Series principal publisher, in 2003. This present volume is published by BAR Publishing, 2019.

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TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES LIST OF PLATES ACKNOWLEDGMENTS ABSTRACT FOREWORD

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CHAPTER ONE. INTRODUCTION AND METHODS Introduction ........... ............. ......................................................... ........... ...................... .................. ................ 1 Brief Historical Perspective ................................................................................................. .......................... 1 Tempo and Mode - Phyletic Gradualism vs. Punctuated Equilibrium ......................................................... 5 Morphological Relationships - Cladisitics vs. Phenetics ....... .................... ................................................... 7 Methods and Data .......... ............. ............................................................................... ........... ....................... 12

CHAPTER TWO. SUB-SAHARAN AFRICA Introduction ............................ ................. .............. ................ ........... ...................................... ...................... 23 Bodo D' Ar ............................ ....................................................... ...................... ............. ... :......................... 23 Ndutu .................................... .................. ................ ..................................................................................... 27 Saldanha-Elandsfontein-Hopefield ....................................................................................... ............. .......... 29 Kabwe-Broken Hill-Rhodesian Man ..................................................... ...................................................... 30 Eyasi ................................ ............................... .......... .................................... ............ ................................... 31 Omo (Kibish) ................... ........................................................................ ......................... ........................... 32 Floris bad .................................................................................................................... ............ ...................... 35 Ileret ......................................................................................................................................... .................... 37 Herto ..................................... ....................................................................................... ................................ 37 Laetoli-Laetolil (Ngaloba Beds) .................................................................. ........................... ..................... 38 Eliye Springs ........................... ....... .................. ................. ........................................ ........... ...................... .. 39 Singa ............. ......... ................................................................................................................. ......... ............ 40 Border .Cave ................................................................................................................................................. 44 Klasies River Mouth ............................................. ...................................................... ·............ ................... .. 4 7 Discussion ..................................................................... ............................................................................... 52 Conclusion ........ ................................................................. ........ ....................................................... ........... 59

CHAPTER THREE. NORTH AFRICA Introduction ......................................................................... ................................................................. ........ 61 Sale .............................................................. ................................................................................................ 61 Mifsud-Giudice Quarry, Kebibat - Rabat .................................................................................................... 61 Jebel lrhoud-Djebel lghoud ......................................................................................................................... 61 Haua Fteah ..................... .............................................................. ................................................... ............. 66 Mugharet El-'Aliya-Tangier (High Cave) ................................. ................................................................. .. 66 Temara ...................... ............... ......................................................... ........................................................... 67

Dar-Es-Soltan(e) ..................................................................................... ..................................................... 67 Discussion ..................................................................................................... ............................................... 68 Lithic Material .............................................................................................................................................. 69 Conclusion ................................................................................................................................................... 70

CHAPTER FOUR. WESTERN ASIA Introduction ............... .................................................................................................... ............................... 71 Dmanisi .............................................. .................................................................... ........................ .............. 71 The Levant ................................................................................................................................................... 72 Galilee-Mugharet el-Zuttiyeh ...................................................................................................................... 72 Mugharet es-Skhul ....................................................................................................................................... 75 J ebel Qafzeh-J ebel Kafzeh .......................................................................................................................... 80 Mugharet et-Tabfin .................................... ................. .................................................... ...................... ....... 84 Amud ............ ............................................. ................. ................................................................................. 89 Mugharet El-Kebarah-Kebara ..................................................................................................................... 91 Dederiyeh ..................................................................................................................................................... 92 Shanidar ........................ ............................................................................................................................... 93 Central Asia ................................................................................................................................................. 97 Mezmaiskaya Cave ..................................... ....................... .......................................................................... 97 Teshik-Tash ........................................................................................................................................ :........ 98 Starosel'e-Staroselye-Staroselje .......................................... ......................................................................... 99 South Asia .................................................................................................................................................. 100 Narmada ..................................................................................................................................................... 100 Discussion ............................. ................................................. .................................................................... 102 Lithic Material ........................................................................................................................................... 105 Conclusion ............. .......... ....................................... ........................ ........................................................... 106

CHAPTER FIVE. EUROPE Introduction ................................................................ ............................................................... ................. 110 Middle Pleistocene Hominids .................................................................................................................... 115 V enta Micena-Orce and Cueva Victoria .............................. .................................................................. .... 117 Ceprano ...................................................................................................................................................... 117 Atapuerca - Gran Dolina ............................................................................................................................ 118 Petralona ...................................................................... ......................... ............................. ........................ 118 Mauer-Heidelberg ................. ...................... ........................................................... ................... ................. 122 V ertesszollos ......... ........................................................ .................................... .................... ..................... 123 Caune de L'Arago-Tautavel ...................................................................... ................................................. 124 Bilzingsleben .......................... ............... .......................................................... .......................... ................ 126 Montm~urin .......................................................................... ......................................... ............................ 127 Steinheim ................................................................................................................................................... 128 Swanscombe ................. ....................................... ...................................................................................... 129 Atapuerca - Sima de los Heusos ................................................................................................................ 132 Biache-Saint-Vaast ................................ ...................................... ................................ .............................. 133 La Chaise-L'abri Suard .............................................................................................................................. 134 F ontechevade ............................................................................................................................................. 134 Ehringsdorf .................................................................................................................................. .............. 135 Reilingen ................................................ ....................................................... ............................................. 137 Upper Pleistocene Hominids ..................................................................................................................... 137 Alicante .......................................................................... ............................................ ................................ 137 Krapina ...................................................................................................................................................... 138 Saccopastore .......................... ..................................................... ............................................................... 142 "Classic" Neanderthals ................................................... ........................................................................... 144 11

Vindija ....................................................................................................................................................... Saint-Cesaire ........................................................................ ...................................................................... Anatomically Modem Upper Palaeolithic ................................................................................................. Discussion .................................................................................................................................................. The Replacement Hypothesis and the Multiregional Hypothesis .............................................................. The Neanderthals: Where From and Why? ............................................................................................... Conclusion ................................................................................................................................................. The Earliest Anatomically Modern Humans In Europe ............................................................................ Lithic Material ........................................................................................................................................... Coexistence and Replacement .......................... ......................................................................................... The Neanderthals - Up Close and Personal ...............................................................................................

148 150 151 151 152 154 161 161 164 167 168

CHAPTER SIX. AUSTRALASIA Introduction ................................................................................................................................................ 171 Island Southeast Asia ................................................................................................................................. 171 Indonesian Homo erectus .......................................................................................................................... 171 The Deep Skull of Niah ............................................................................................................................. 178 Tabon Cave ................................................................................................................................................ 179 Australia ........................................................................................................................................ ............. 181 Lake Mungo ............................................................................................................................................... 181 Lake Garnpung - WLH 50 ........................................................................................................................ 185 Discussion .................................................................................................................................................. 187 The Replacement Hypothesis .................................................................................................................... 187 The Regional Continuity or Multiregional Hypothesis ............................................................................. 188 Regional Features ...................................................................................................................................... 189 Conclusion ................................................................................................................................................. 211

CHAPTER SEVEN. EAST ASIA Introduction ................................................................................................................................................ 214 Homo erectus ............................................................................................................................................. 214 Longgupo ................................................................................................................................................... 214 Lan ti an ....................................................................................................................................................... 215 Yuanmou .................................................................................................................................................... 216 Yunxian ............................................................................................................................. ......................... 216 Zhoukoudien (Choukoutien) Locality 1 .................................................................................................... 216 Nanjing ...................................................................................................................................................... 217 Hexian ........................................................................................................................................................ 218 Archaic Homo sapiens ............................................................................................................................... 218 Jinniushan ...................................................................................................................... ............................ 218 Dali ............................................................................................................................................................ 219 Maba-Mapa .......................................................................................................................................... ...... 220 Xujiayao ..................................................................................................................................................... 222 Early Homo sapiens ................................................................................................................................... 223 Liujiang ...................................................................................................................................................... 223 Zhoukoudian Upper Cave .......................................................................................................................... 224 Minatogawa ............................................................................................................................................... 227 Discussion .................................................................................................................................................. 229 The Replacement Hypothesis .......................... .......................................................................................... 229 The Regional Continuity or Multiregional Hypothesis ................................. ............................................ 229 Regional Feature ........................................................................................................................................ 231 Lithic Material ........................................................................................................................................... 240 Conclusion ................................................................................................................................................. 241

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CHAPTER EIGHT. CONCLUSION Introduction ............................................................................................................... ................................. 245 The Regional Continuity or Multiregional Hypothesis ............................................................................. 245 The Single Origin and Replacement Hypothesis .............................................................. ......................... 247 Genetic Data .............................................................................................................................................. 247 Prehistoric mtDNA .................................................................................................................................... 253 Discussion .................................................................................................................................. ................ 254 Sub-Saharan Africa .................................................................................................................................... 254 North Africa .................................................................. ...................... ....................................................... 254 Western Asia .............................................................................................................................................. 255 Europe ........................................................................................................................................................ 256 Australasia ................................................................................................................................................. 257 East Asia ......................................................... ........................................................... ................................ 25 8 Conclusion ................................................................................................................................................. 260

BIBLIOGRAPHY

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APPENDIX TABLES

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PLATES

LIST OF TABLES 1: 1 Cranial measurements used in analyses, their abbreviations and sources where they are defined. 1:2 Crania that formed the basic data set for all metrical analyses and abbreviations for Figures and Tables. 1:3 Discontinuous, discrete, epigenetic, or non-metric traits used in analyses. 1:4 Discontinuous, discrete, epigenetic, or non-metric mandibular traits used in this study. 2: 1 Sub-Saharan African crania examined, abbreviations for Figures and Tables, and source of data. 2:2 K-means cluster analysis with Ndutu. 2:3 K-means cluster analysis with Singa, Omo 1, Omo 2 and Saldanha 1. 2:4 K-means cluster analysis with Laetoli H18, Omo 2, Saccopastore 1 and Petralona. 2:5 K-means cluster analysis with Eliye Springs KNM-ES 11693. 2:6 K-means cluster analysis with Singa. 3: 1 North African hominids examined, abbreviations for Figures and Tables, and source of data. 3:32 K-means cluster analysis with Jebel Irhoud 1. 4: 1 Wes tern Asian hominids examined, abbreviations for Figures and Tables, and source of data. 4:2 K-means cluster analysis with Skhiil 4, Skhul 9, Jebel Qafzeh 6, Jebel Qafzeh 9, and Jebel Irhoud 1. 4:3 K-means cluster analysis with Skhul 4, Jebel Irhoud 1 and Jebel Qafzeh 6, 9 and 11. 4:4 K-means cluster analysis with Tabun Cl, Shanidar 1 and Amud 1. 4:5 K-means cluster analysis with Shanidar 5. 4:6 Measurements used in the Ordered Similarity Matrix for the Levant with Jebel lrhoud 1. 4:7 Ordered Similarity Matrix for the Levant with Jebel Irhoud 1. 4:8 Brachia! and crural indices for Western Asian hominids. 5: 1 Correlation chart for the European Middle and Upper Pleistocene. 5:2 European crania examined, abbreviations for Figures and Tables, and source of data. 5:3 K-means cluster analysis with Saccopastore 1 and Petralona. 5:4 K-means cluster analysis of European core group . 5:5 K-means cluster analysis with Guattari-Monte Circeo 1, Saccopastore 1, Krapina C, La Quina H18 and Teshik-Tash. 5:6 Some characteristic features of the cranium and mandible evident on the Neanderthals. 5:7 Measurements used in the Ordered Similarity Matrix for Europe. 5: 8 Ordered Similarity Matrix for Europe. 6: 1 Australasian crania examined , abbreviations for Figures and Tables, and source of the data. 6:2 Measurements used in the Ordered Similarity Matrix for Australasia. 6:3 Ordered Similarity Matrix for Australasia. 6.4 K-means cluster analysis with some of the Ngandong crania. 6:5 The occurrence of"Regional Continuity Features" . 6:6 Percentages of the occurrence of proposed "Regional Features" on Australian Aboriginal crania. 7: 1 East Asian crania examined, abbreviations for Figures and Tables, and source of data. 7:2 K-means cluster analysis with Dali cranium. 7:3 The occurrence of "Regional Continuity Features" for East Asia. 7:4 Regional variation of three dental traits used as Regional Continuity Features. 7:5 Comparison of facial morphology on early Homo sapiens material from east Asia and Generalised Mongoloid form. 8: 1 Late Pleistocene and early to mid-Holocene core group of crania studied. 8:2 Cranial measurements used in analyses. 8:3 Hominid crania compared to Late Pleistocene and early to mid-Holocene core group. 8:4 Results ofk-means cluster analyses.

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14 15 17 18 24 27 29 33 40 42 62 63 71 77 82 87 95 107 107 108 111 112 122 129 140 145 162 162 171 177 177 178 193 194 214 220 235 237 242 261 261 262 262

LIST OF FIGURES 1: 1 Cranial non-metric traits used in this study. 16 1:2 Mandibular non-metric traits used in this study . 18 2: 1 Map showing Sub-Saharan African sites mentioned in text. 25 2:2 The chronological position of the Sub-Saharan African hominid sample . 26 2:3 Dendrogram from sum of squares cluster analysis with the Ndutu cranium . 28 2:4 Correspondence analysis with Ndutu. 28 2:5 Dendrogram from sum of squares cluster analysis with Omo 2 and Laetoli H18. 34 2:6 Correspondence analysis with Omo 2 and Laetoli H18. 34 2:7 Dendrogram from sum of squares cluster analysis with Eliye Springs 11693. 41 2:8 Correspondence analysis with Eliye Springs 11693. 41 2:9 Dendrogram from sum of squares cluster analysis with the Singa calvaria. 43 2: 10 Correspondence analysis with the Singa calvaria . 43 3: 1 Map showing North African sites mentioned in text. 62 3 :2 Dendrogram from sum of squares cluster analysis with Jebel Irhoud 1. 64 3 :3 Correspondence analysis with Jebel Irhoud 1. 64 4: 1 Map showing the Levantine sites mentioned in the text. 72 4:2 The probable chronological position of the Western Asian hominid sample. 73 4:3 Dendrogram from sum of squares cluster analysis with Skhul 4, 5 and 9, Jebel Qafzeh 6 and 9 and Jebel Irhoud 1. 78 4:4 Correspondence analysis with Skhul 4, 5 and 9, Jebel Qafzeh 6 and 9 and Jebel lrhoud 1. 78 4:5 Dendrogram from sum of squares cluster analysis with Skhiil 4 and 5, Jebel Irhoud 1 and Jebel Qafzeh 6, 9 and 11. 83 4:6 Correspondence analysis with Skhul 4 and 5, Jebel Irhoud 1 and Jebel Qafzeh 6, 9 and 11. 83 4:7 Dendrogram from sum of squares cluster analysis with Tabiin Cl, Amud 1 and Shanidar 1. 88 4:8 Correspondence analysis with Tabun Cl, Amud 1 and Shanidar 1. 88 4:9 Map showing western and central Asian sites mentioned in text. 93 4 :·10 Dendrogram from sum of squares cluster analysis with Shanidar 5. 96 4: 11 Correspondence analysis with Shanidar 5. 96 4: 12 Map showing Indian sites mentioned in text. 101 4: 13 Correspondence analysis with Narmada and Homo erectus material. 101 5: 1 Map showing some European sites mentioned in text. 110 5:2 Map showing additional French sites mentioned in text. 113 5:3 Map showing German sites mentioned in text. 114 5:4 Map showing Czech Republic , Slovak Republic and Austrian sites mentioned in text. 114 5:5 Map showing Hungarian and Croatian sites mentioned in text. 115 5:6 The probable chronological position of the European hominid sample. 116 5:7 Dendrogram from sum of squares cluster analysis with Petralona and Saccopastore 1. 121 5:8 Correspondence analysis with Petralona and Saccopastore 1. 121 5:9 Scatter diagram from principal components analysis of core group of crania. 130 5: 10 Dendrogram from sum of squares cluster analysis of core group of crania. 130 5: 11 Correspondence analysis of core group of crania. 131 5:12 Dendrogram of sum of squares cluster analysis with Guattari-Monte Circeo 1, Krapina C, La Quina Hl8 , Teshik-Tash and Saccopastore 1. 141 5:13 Correspondence analysis with Guattari-Monte Circeo 1, K.rapina C, La Quina H18, Teshik-Tash and Saccopastore 1. 141 5:14 Correspondence analysis of the occipital squama. 143 5: 15 Correspondence analysis of cranial non-metric traits . 146 5: 16 Correspondence analysis of mandibular non-metric traits. 146 6: 1 Map showing Southeast Asian sites mentioned in the text. 172 6:2 Dendrogram from sum of squares cluster analysis with calvaria from Ngandong . 178 6:3 Map showing Australian sites mentioned in the text. 182 7: 1 Map showing Chinese sites mentioned in the text. 215 7 :2 Dendrogram from sum of squares cluster analysis with the Dali cranium. 221 7 :3 Correspondence analysis with the Dali cranium. 221 7:4. The chronological position of the East Asian hominid sample. 228 8: 1 Map showing early archaic Homo sapiens material that displays grade similarities . 255 8:2 Dendrogram from group average cluster analysis of Late Pleistocene and early to mid-Holocene core group. 263 8:3 Dendrogram from group average cluster analysis of core group with Skhiil 4 & 5, Jebel Qafzeh 6 & Jebel Irhoud 1. 265 8:4 Dendrogram from group average cluster analysis of core group with European Upper Palaeolithic crania. 265 8:5 Dual Source Model. 268 8:6 Assimilation and Replacement Model. 269

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LIST OF PLATES Plate 1. (casts) Top: Bodo (left) and Petralona (right) Bottom:Petralona (left) and Bodo (right) Plate 2. (cast) Laetoli H 18 Plate 3. (casts) Top: Florisbad (old reconstruction) Bottom: Jebel Irhoud I Plate 4. (casts) Amud I (left) and Skhul 5 (right) Plate 5. (casts) Top and Middle: Amud I (left) and Jebel Qafzeh 9 (right) Bottom: Tabun Cl (left) and Jebel Qafzeh 9 (right) Plate 6. (casts) Skhul 5 (left) and Jebel Qafzeh 9 (right) Plate 7. (casts) Amud I (left) and La Ferrassie 1 (right) Plate 8. (casts) Top: Mandibles of Skhul 5, Amud I and Jebel Qafzeh 9 Bottom: Petralona (left) and Arago 21 (right) Plate 9. (cast) Arago reconstruction, comprising Arago 21 and Arago 47 Plate 10. (casts) Top: La Ferrassie I (left) and Krapina C (right) Bottom: Gibraltar I (left) and Krapina C (right) Plate 11. (casts) Top: Gibraltar l(left) and La Ferrassie I (right) Bottom: La Quina HS (left) and Gibraltar 1 (right) Plate 12. (cast) Saint-Cesaire Plate 13. (cast) La Ferrassie I (left) and Cro-Magnon I (right) Plate 14. (casts) Top: Sangiran 17 (left) and Kow Swamp I (right) Middle: Lake Mungo 3 Bottom: Kow Swamp 15 (left) and Kabwe I (right) Plate 15. (casts) Close-up of the splanchnocranium Top: Kow Swamp 15 Bottom: Kow Swamp 15 (left) and Sangiran 17 (right)

ACKNOWLEDGEMENTS Antiquities Nationales de Saint Germain-en-Laye; Laboratoire d 'Anthropologie, Universite de Bordeaux; Institut Royal des Sciences Naturelles des Belgique, Brussels; Laboratoire de Paleontologie animale, Universite de Liege; Abteilung Anthropologie, Naturhistorisches Museum Wien; Rheinisches Landesmuseum, Bonn; Staatliches Museum fiir Naturkunde, Stuttgart; Geologisch-Palaontologisches Institut der Universitat Heidelberg; Institut fiir Humanbiologie, Universitat Hamburg; Museo di Antropologia, Dipartimento di Biologia Animale e Dell 'Uomo , Universita di Roma ; Instituto de Paleontologia de Sabadell; Caledratico de Antropologia Fisica, Universidad de Barcelona; Museo Prehistoria, Valencia; Zarod za Paleontologiju i Geologiju uvartora , Zagreb; Geolosko-Paleontoloski Muzej, Zagreb; Department of Anthropology , Harvard University; Department of Anatomy, University of the Witwatersrand; National Museum, Bloemfontein; South African Museum , Cape Town; Department of Archaeology, University of Stellenbosch; Department of Anatomy, Medical School, University of Cape Town; Rockerfeller Museum, Jerusalem; Department of Prehistory, Research School of Pacific Studies, Australian National University, Canberra; Department of Anthropology, Western Australian Museum.

This volume is an amalgamation of research undertaken over many years and that relied on the generosity of many colleagues who have shared their knowledge and/or given me access to material under their care. I would especially like to thank Dr. M.J. Walker for his friendship, guidance and support over many years. Professor R.V.S. Wright provided essential advice and help with the computer programs used and the interpretation of the kilometres of computer printout. I am also especially grateful to Professor C.B. Stringer who provided crucial data, advice and direction to my research , as well as friendship and hospitality during my visits to the British Museum (Natural History) . During my world odysseys to study hominids, visit sites and participate in fieldwork I was fortunate enough to meet many palaeontologists and archaeologists who were most willing to share their vast amounts of knowledge on human evolutionary history. Those I would especially like to thank are Professor F .H. Smith, Professor H.J . Deacon , Professor M.H . Wolpoff, Dr. F. Spencer, Professor G.P. Rightmire, Dr. G. Brauer and Professor C. Gamble. In Australia I would like to thank Professor R.V .S. Wright, Dr. R. Fletcher, Dr. J.P. White, Dr. S.J. Bourke, Professor A.G. Thome, Dr. C. Pardoe and Professor C.P. Groves. I am also in debt to Dr. E. Delson, Dr. I Tattersall, Professor P.V. Tobias, Professor P. Mellars and Professor C.B. Stringer for allowing me to attend and participate in some important gatherings on human evolution when I was undertaking my Doctoral dissertation, at which I was able to examine and discuss hominid skeletal morphology with colleagues from around the world. The influence on my thinking that these occasions had was substantial.

The Carlyle Greenwell Research Fund, University of Sydney and the Australian Institute of Aboriginal Studies provided financial support for some of the research upon which this volume is based.

I would like to acknowledge my gratitude and indebtedness to the following institutions for permitting me to examine hominid material in their collections: British Museum (Natural History) London; Musee de I'Homme and lnstitut de Paleontologie Humaine, Paris; Laboratoire de Prehistoire du Museum National D'Histoire Naturelle; Laboratoire de Paleontologie Humaine et de Prehistoire , Universite de Provence , Marseilles ; Laboratoire de Paleontologie des Vertebres et Paleon!ologie Humaine, Universite de Paris VI; Musee des

Finally, Dr. Natalie Franklin kindly produced the diagrams and provided support during the researching and writing of this work and so to her I owe a great deal.

Assistance in data collection was provided by Dr. Stephen Bourke, Ms Catriona Bonfiglioli, Dr. Mike Roland and Dr. Robert Kruszynski, which was much appreciated.

Phillip J Habgood 2003

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ABSTRACT It is established that the characters proposed to be "regional continuity traits" linking Homo erectus material with late

There are two major explanations for the world-wide ongm of anatomically modem humans: the Replacement Hypothesis and the Multiregional Hypothesis. The Replacement Hypothesis proposes that anatomically modem Homo sapiens evolved in one main area from where they radiated out and replaced the local populations in all other parts of the Old World. This explanation assumes that there was a relatively recent common Homo sapiens ancestral population. The area where anatomically modem humans originally developed is generally given as sub-Saharan Africa, although western Asia also remains a possibility. The Multiregional Hypothesis emphasises regional morphological continuity and argues that anatomically modem humans developed from morphologically separate, although not genetically isolated, groups (Homo erectusarchaic Homo sapiens) that had already differentiated from each other. The Multiregional Hypothesis contends that regional variations have developed at the species' peripheries and are maintained by a balance between gene flow and selection, which caused the formation of morphological clines. Inherent within this explanation is gene flow across the species' distribution and it is the balance between selection, gene flow and genetic drift that underpins the Multiregional Hypothesis .

Pleistocene and early Holocene crania and modem populations in Australasia and east Asia are retained primitive features present on Homo erectus and archaic Homo sapiens crania and mandibles in general. Many are also commonly found on the crania and mandibles of anatomically modem Homo sapiens from other geographical regions. However, there are a number of morphological features which, when found in combination, appear to document a degree of morphological continuity between the archaic hominids and later early modem material. For Australasia the combined features are - a long and sagittally flat frontal bone, posteriorly placed minimum frontal breadth, a prognathic face and malars with everted lower borders and prominent malar tuberosities. For east Asia the combined features, which may be related to facial flatness, are - more perpendicularly orientated nasal bones, frontonasal and frontomaxillary sutures on almost the same level, a non-depressed nasion, an angular rather than a rounded junction of the zygomatic bone and the zygomatic process of the maxilla or presence of an incisura malaris and possibly also shovel-shaped upper incisors. One could argue that there are no compelling reasons to reject the Multiregional explanation for the evolution of anatomically modem humans in Australasia and east Asia. There are, however, problems with this argument in that some of the early modem crania from these regions do not have the morphological pattern that would be expected if there had been continuity from Homo erectus through to modem Homo sapiens.

The problem of the origin of anatomically modem Homo sapiens throughout the Old World has been addressed in this study. The results from both multivariate analyses of cranial and mandibular metric and/or non-metric data and morphological studies are used to investigate when anatomically modem humans first appeared in sub-Saharan Africa, north Africa, western Asia, Europe, Australasia and east Asia, so as to ascertain which of the two explanations best explain the world-wide appearance of anatomically modem Homo sapiens.

Neither of the two explanations (Replacement or Multiregional Hypotheses) can explain the appearance of anatomically modem humans throughout the entire Old World. In some areas there is clear evidence for continuity while in other areas the evidence is inconclusive, or indicates replacement.

It was found that anatomically modem Homo sapiens appears to have developed in situ from archaic Homo sapiens in sub-Saharan Africa, north Africa and western Asia. Whilst there are issues with the dating of most of the "transitional" specimens, the best estimate of the date at which the earliest anatomically modem humans first appeared in these regions is approximately 150-100,000 years BP, suggested by the sites of Herto, Klasies River Mouth, Jebel Irhoud, Skhul and Jebel Qafzeh. However, it is not known if anatomically modem humans evolved first in one of these areas and then influenced the course of evolution in the other areas (i.e. gene flow and/or hybridisation).

On the current evidence one could even argue that there were two large centres of origin for anatomically modem Homo sapiens : one in the western sector of the Old World - Africa and western Asia, and the other in the eastern sector of the Old World - east Asia and Australasia. However, these two regions would not have been independent of each other, but rather there would have been multidirectional gene flow and migration between them throughout much of the Middle and Upper Pleistocene. Europe is the only area that clearly documents the migration of anatomically modem humans into the region .

The hominid sample from Europe presents the gradual development of the Neanderthal morphological pattern over a 250,000 year period . Europe does not present any conclusive evidence for the local in situ evolution of anatomically modem Homo sapiens out of the Neanderthals, but rather documents the replacement of the Neanderthals by modem humans and the replacement of Middle Palaeolithic artefact assemblages by Upper Palaeolithic assemblages. These changes are especially evident in western Europe . Mitochondrial DNA (mtDNA) recovered from a number of Neanderthal and anatomically modem Upper Palaeolithic remains has been interpreted as indicating that the Neanderthals did not contribute to the mtDNA pool of modem humans and therefore supports the Replacement Hypothesis . The mtDNA results would seem to exclude any significant interbreeding between Neanderthals and anatomically modem Upper Palaeolithic humans.

However, the morphological, archaeological and genetic data provides a complex image of the transition from archaic to anatomically modem Homo sapiens across the world that necessitates an explanation that combines some local continuity, some population replacement, and significant multidirectional gene flow. This complex phenomenon can best be described as an "Assimilation and Replacement" model, where Europe saw the replacement of the Neanderthals by anatomically modem Homo sapiens, whereas eastern Asia experienced the assimilation of anatomically modem humans and the archaic indigenous populations. However, at the present time it is not possible to conclusively establish the relative importance of replacement and continuity in 'peripheral' areas such as east Asia and southeast Asia.

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FOREWORD evolutionary biology (though it should be mentioned here that the palaeoprimatologist Fred Szalay has argued cogently1 that cladistical phylogenetic analysis by no means fulfils all of the criteria necessary for a correct application of that approach, notwithstanding the profession of adhesion to it by no less than the founder of phylogenetic analysis himself, Willi Hennig).

Phil Habgood is that valued friend and scholar we all hope will be a felicitous spin-off when we give both encouragement and free rein to an outstanding doctoral candidate. This volume is a much revised version of the thesis for which he was awarded the Ph.D. in 1988. His work stands here on its own merits. Its importance lies in the way in which he applies numerical taxonomy of Pleistocene human skulls to guard against accepting extreme positions in interpreting human evolution .

Mathematical representation in science is often a hall-mark of sceptical scientific critical realism because it is the key that opens up a gateway to counter-intuitive explanation based on how data behave, capable of drawing contrasts between conflicting claims about reality. This is no less true in evolutionary biology than in cosmogeny. Phil Habgood shows how numerical taxonomy applied to Pleistocene skull measurements can give results that are not only in line with those produced by other methods, but have an undoubted advantage over them of being backed up by rigorous methodology. They highlight serious difficulties with a Multiregional interpretation of Pleistocene human evolution, while indicating some possible drawbacks to an interpretation of modem human origins based wholly on a total Replacement (Out-of-Africa) hypothesis.

Etched in my memory from 1960 is the Professor of Anatomy, Sir Wilfred Le Gros Clark, announcing to us preclinical medical students at Oxford that "Science is measurement, ladies and gentlemen, science is measurement!" His wise dictum underpins the significance of numerical considerations in taxonomy. This significance has three epistemologically important aspects, each of which distinguishes the modem natural scientist's approach to natural phenomena from the traditional naturalist's dilettantism informed by deductive arguments chosen all too often in self-serving fashion. First, measurements imply an approach to inquiry through fragmentation, which is incompatible with a total, holistic appreciation of the individual objects of our attention, no matter how dispassionately and completely they be observed, or comprehensively described, whether as individuals or in wider context. Secondly, measurements belong to a particular class of observations that is distinguished by our ability to transform them, usually by mathematics, into data that are abstractions, and to that extent unreal, though by no means unrealistic. Thirdly, our selection of both a raw measurement and its transformation as a datum implies an interdependence between them which may be inferred methodologically or theoretically, but which must ultimately be held up to reality with a critical and skeptical attitude that is prepared to admit a possibility that new data may not always be readily accommodated within known patterns, and may not so much corroborate accepted models as require adjustments to them.

Phil Habgood's minimalist conclusion is that our own ancestral stock achieved widespread replacement of earlier human groups though some of the latter may very occasionally have undergone assimilation. His proposal combines what Chris Stringer2 considers to be two options situated between the extreme interpretations of total Replacement out of Africa and Multiregional evolution, namely, the possibilities of African Hybridization with Replacement and Assimilation. These two options share much in common, and are now receiving more consideration than either of the embattled stances of the warring ideologues of Multiregional and total Replacement interpretations. As I have remarked elsewhere, "the battle has nowadays been reduced to skirmishes in the middle ground" 3• Phil Habgood was trained as both an archaeologist and anthropologist in the Arts Faculty at Sydney University (where I taught between 1973 and 1988). This volume speaks loudly for his ability to correct, with a dose of scientific critical realism, the pervasive influence of so many Arts Faculty teachers who interpret human thought, behaviour, society, and history, in discourses that are cloaked in mantles of academic respectability with common threads of all too complicated philosophical arguments informed by an all too uncomplicated intuitive realism. However unsurprising it may be that their discourses are usually incommensurable with those to which belong such scientifically-inferred invisible realities as are quarks, black holes, prions, or genes, it is undeniable that their "what-yousee-is-what-you-get" intuition of reality often immunizes their discourses, in self-serving manner, against attempts to subvert them, because the notion of reality on which they are based is fundamentally incommensurable with a sceptically critical one.

The scientist's critical realism leads to easy acceptance of counter-intuitive notions (e.g. biological speciation by natural selection working through populations rather than individuals; concepts in matrix algebra such as the square root of ~nus one and hyperspace on which principal component analysis depends), and even to acceptance of the usefulness, according to the matter in hand, of alternative counter-intuitive concepts that seem ostensively antagonistic (e.g. a particle theory alongside a wave theory of light; quantum mechanics alongside general relativity; a biological species-concept appealing to observable reproductive processes, an ecological species-concept appealing to separation and separability of habitats, a mate-recognition species-concept appealing to ethological observation, and morphologically-based species-concepts - the only ones available for classifying extinct fossils - such as the inclusive evolutionary phyletic species-concept, and the exclusively cladistic, or phylogenetic, species-concept). Sceptical scientific critical realism is the appropriate approach to inquiry in all branche s of modem science, including

This can cause severe epistemological headaches in archaeology and anthropology when quasi-ethnographical, X

conceptions of reality (not to mention relativist and swprise to most natural scientists, who tend to be sceptical, critical and cautious, even if the bleakness of their grudging conclusions irks those archaeologists, anthropologists, and evolutionary biologists, who strive to find a short cut to understanding the past in terms of perspectives from the present - not to mention those optimistic and immodest phylogenetic ideologues among them who assert with hubris to be already taking it.

quasi-historical, or even quasi-biological, positivist postmodernist ones), and theoretical deductions therefrom, are deployed to interpret material phenomena, be they flints or skulls: namely, should "awkward" phenomena be explained away (e.g. as "exceptions which prove the rule", or as unusual cases to be filed in the "pending" tray), or should they be held up so as to question the veracity of the preferred interpretation imposed on them and thus demand methodological or theoretical adjustments to it? Scientific sceptical criticism might prefer -as Phil Habgood does hereto circumscribe the scope of inquiry to only that universe of commensurable phenomena to which a prescriptive methodology, of observations and data on them (fossil skulls), offers a normative epistemological foundation for determining just what can be known, however little it be, and just how far the frontiers of knowledge can thereby be pushed back, if barely at all, thus reducing, however slightly, the extent of our ignorance.

Phil Habgood's dual training in archaeology and anthropology confers a singular advantage when it comes to assessing the contexts in which different fossil hominids have been found. First-hand experience of excavating archaeological sites helps us better to interpret reports about where, how, and when human fossils have been found - as well as how to read between their lines, and perhaps to wonder about what might even have been left out of some accounts. Phil Habgood has travelled around the world to get first-hand observations, measurements, and data, wherever possible on original fossil specimens, and by visiting sites where they were found. His is very far from being a study based merely on casts of fossils, second-hand sources, and rehashed accounts of what earlier generations had recorded. It is therefore not an easy read. All the same, it should be required reading for all who have a serious interest both in human origins themselves, and in how we know what we can know about them to the best of our ability.

Quasi-biological discourses (e.g. "evolutionary psychology", etc.) give particular grounds for concern because they claim to be justified by the results of inquiry in the natural sciences, when perhaps the most that a critical sceptic might allow is that they need not necessarily be in conflict with some of those results. They may be forgiven to the extent that some natural scientists themselves have stumbled into epistemological quagmires, as already remarked in connexion with Willi Hennig. They might also be forgiven to the extent that quasi-evolutionary arguments, couched in that rhetorical descriptive language of nineteenth-century naturalists imbued with intuitive realism (superseded already in the mid-twentieth century by the unrhetorical prosaic scientific biological language of the Neo-Darwinian evolutionary synthesis with modem genetics) which so appeals to the cladistic naturalist's simple-mindedly uncomplicated intuitive realities of fossil hominid phylogenetic species, are arguments that make up a far easier stick with which to beat Creationists in public than do the conclusions of Phil Habgood's multivariate statistics, deployed here in the form of principal component analysis where counter-intuitive reality, however much it depends on the unreal numbers that underpin the mathematics of matrix algebra and the multidimensionality of hyperspace, is not at all unrealistic from the standpoint of the sceptical critical realism of the natural scientist.

Michael J. Walker Professor of Physical Anthropology Area de Antropologia Fisica Departamento de Zoologia y Antropologia Fisica Facultad de Biologia Universidad de Murcia 30100 Murcia, SPAIN http://www.um.es/antropfisica 1 Szalay, F. (1993) Species concepts. The tested, the untestable, and the redundant. In, Species, species concepts, and Primate evolution. W. H. Kimbel and L. N. Martin, eds, pp. 21-41. "Advances in primatology", Plenum Press, New York and London. 2 Stringer, C. B. (2002) Modem human origins: progress and prospects. Philosophical Transactions of the Royal Society of London, series B, 357: 563-579. 3 Walker, M. J. (2003) The quest for our human ancestors. The Review of Archaeology 24: 20-38.

Phil Habgood's approach is therefore much to be preferred, even though, by following it, the boundaries between different species-concepts (see above) will long remain fuzzy when particular groupings of fossil skulls are considered, let alone individual skulls. That, however, will come as no

XI

CHAPTER ONE. INTRODUCTION AND METHODS Introduction Pilbeam has suggested that The evolutionary transition to modem humans is one of the most dramatic of all steps in hominid evolution, and it is the only one that is reasonably well documented both anatomically and archaeologically. (1986:337) The problem of the origin of anatomically modem Homo sapiens throughout the Old World is being addressed in this study. Multivariate analyses and morphological comparisons are used to investigate when anatomically modem humans first appeared in sub-Saharan Africa, north Africa, western Asia, Europe, Australasia and east Asia in order to evaluate the major explanations for the world-wide appearance of anatomically modem Homo sapiens.

Brief Historical Perspective As has been pointed out by many authors the interpretation of the hominid fossil record has been (and still is being) greatly influenced by the intellectual predilections of society at the time (e.g. Boaz 1982; Brace 1964, 1981; Daniel 1967, 1978; Eiseley 1957; Haddon 1910; Hammond 1979, 1980, 1982; Hooton 1931; Millar 1974; Oakley 1976; Spencer 1984; Spencer and Smith 1981; Smith 1987; Trinkaus 1982d; Weiner 1955). This is especially so with those hominids that directly preceded the appearance of anatomically modem humans. As pointed out by Constable (1973), archaic Homo sapiens (especially Neanderthals which will, following Howells 1974a and contra Vallois 1952, be spelt with an "h") are still often viewed as shambling, beetle-browed louts that prowled the earth wearing ragged furs, speaking in gru~ts and occasionally pausing to hit women over the head with clubs and drag them back to their caves. It is not until the appearance of anatomically modem Homo sapiens, who essentially looked like modem peoples, that some "nobility" was bestowed upon prehistoric humans. Society's predilections and prejudices have, therefore, influenced the interpretation of when and where anatomically modem humans originated. Human Antiquity The study of prehistoric humans has had a long history (Daniel 1959, 1978; Grayson 1983; Haddon 1910; Hoo!on 1931; Oakley 1964), with major developments occumng during the second half of the nineteenth century. Grayson (1983) documented the gradual recognition of the high antiquity of prehistoric humans, which was a necessary prerequisite for the acceptance of human evolution. Befo~e the publication of Charles Darwin's The Origin of Species m 1859 there were two major competing schools: the "catastrophists" and "transmutationists".

Toe "catastrophists" argued that past life had been destroyed several times ( explaining the appearance of different suites of fossils in different geological strata), as in the Biblical flood, and then created anew. They believed in the fixity of species and that life was the result of miraculous creation. Within

this school of thought were the monogenists who upheld the scriptural unity of all peoples in the single creation of Adam and Eve and thought that modem races had degenerated from Eden's perfection at differing rates, whites the least and blacks the most (Gould 1981). The "transmutationists" rejected creationism, contending that all species were derived from other species by some unknown law. This unknown law was formalised in The Origin of Species (Darwin 1859) in which species were said not to be "fixed", but to have gradually changed. Natural selection was suggested as the mechanism of change that caused the "transmutations" to occur. Within this group would have been the polygenists who abandoned the scriptures and argued that races were separate biological species and so were, as Gould ( 1981) put it, descendants of different Adams. The Neanderthals and Other Hominid Fossils Darwin ( 1859) did not discuss in any detail human evolution in The Origin of Species, but he did deal with it in The Descent of Man published in 1871. One problem Darwin had was a lack of hominid remains. In 1829 and 1830 the remains of four hominids, including the calvaria of an immature Neanderthal, were recovered from a cave near the village of Engis, Belgium (Oakley et al. 1971), but they had little impact on the scientific community of the time. The discovery of a Neanderthal cranium at Forbes' Quarry, Gibraltar, was announced in 1848, but again it went virtually unnoticed.

The discovery of hominid remains in the Feldhofer cave within the Neander Valley, Germany (whence the name Neanderthal is derived), in 1856 (Schaaffhausen 1858) did, however, generate a major scientific debate. Most European scholars regarded these remains as either a modem freak or a member of some savage and barbarous "primitive" race, with one scholar suggesting that the remains belonged to a Mongolian Cossack with advanced rickets (Millar 1974). In Britain, where Darwin's ideas were more widely accepted (Hammond 1982), the Neander Valley remains were placed within an evolutionary framework and generally regarded as a primitive ancestor of modem man. King (1864) assigned the remains to the species "Homo neanderthalensis", highlighting the differences he perceived between them and modem humans. It is highly probable that the previously discovered Neanderthal remains from Eng is and Gibraltar were overlooked because human antiquity had not been clearly established, and because there was no theory of evolution within which they could be placed and which could explain their unusual morphologies. Additional Neanderthal remains were recovered during the latter part of the nineteenth century (Oakley et al. 1971). The recovery of two adult skeletons at Spy, Belgium, in 1886 were the most important of these discoveries in that they confirmed the existence of a distinct fossil type, the Neanderthals.

The remains of anatomically modern humans were also recovered during this period (Oakley et al. 1971). The most famous of these discoveries were the Upper Palaeolithic burials from the Cro-Magnon rockshelter in France. These remains were used by creationists to disprove the theory of evolution because they were said to demonstrate that the earliest human types were the same as modern humans. These remains also opened up the possibility that the Neanderthals, who they appeared to have been contemporary with, were not the ancestors of modern humans.

The Piltdown Men This view was further strengthened with the Piltdown discoveries (announced in Nature on December 5th 1912). "Eoanthropus dawsoni" or "The dawn man of Dawson" consisted of a simian jaw and hominid vault (Dawson and Smith Woodward 1913, 1914). This material displayed a combination of morphological features that had been anticipated by many anatomists, especially Elliot Smith, as a necessary stage in human evolution (Hammond 1979; Miller 1974; Weiner 1955). Additional vault fragments and an isolated molar were also recovered from Piltdown in 1915. Piltdown Man contradicted the "Pithecanthropus erectus" remains from Indonesia, because it indicated that enlargement of the brain occurred before other morphological modifications. The association of the vault and jaw was not accepted by all scholars while some, like Keith ( 1915), who accepted the find as authentic, relegated it, like the Neanderthals, to a side branch and so did not regard it as a direct ancestor of modern humans. Hammond has suggested that ...the historical role of this forgery was not as an actual missing link to man, but as a prehistoric sign-post pointing away from the Pithecanthropine-Neanderthal line towards some other, yet undiscovered, true forefather. (1979:56)

During the 1890s Dubois made discoveries of hominids in Indonesia, which were taxonomically classified as "Pithecanthropus erectus" (Dubois 1896; Oakley et al. 1971). The calotte and the possibly associated femur from Trinil (Day 1984; Day and Mollison 1973) seemed to indicate that bipedalism had preceded the enlargement of the brain. At this time many European scholars regarded these remains as of simian origin, whereas many British workers thought they were human. By the beginning of the twentieth century most palaeontologists envisaged a unilinear phylogenetic line in which human evolution passed from a simian origin to a modern human type (Hammond 1979). Boule was to drastically change this view (Hammond 1979, 1980, 1982).

Eastern European scholars such as Schwalbe, Hrdlicka, Gorjanovic-Kramberger and Weidenreich did not accept Boule's rejection of the Neanderthals as the ancestors of modern humans and they developed what has become known as the Neanderthal Phase of Man Hypothesis in which it was proposed that there was a Neanderthal stage through which hominids had to pass so as to become anatomically modern Homo sapiens (Brace 1981; Hrdlicka 1927, 1930; Radovcic 1985; Spencer 1984; Spencer and Smith 1981; Smith 1987; Weidenreich 1943b, 1947). This difference of views between eastern and western scholars would have been accentuated by the First and Second World Wars. As Hammond (1982) has observed, the French expulsion of the Neanderthals from human ancestry and the British Piltdown finds were allies against the German Neanderthal Phase Hypothesis.

During the first two decades of the twentieth century a number of major discoveries of Neanderthal burials were made in France (Oakley et al. 1971; but see Gargett [1989, 1999] who argued there was no conclusive evidence for Middle Palaeolithic burials in Europe). Boule (1913) used the pathological La Chapelle-aux-Saints skeleton (Straus and Cave 1957; Trinkaus 1985b) for a detailed reconstruction of a "typical" Neanderthal (although he used other Neanderthal remains to reconstruct areas not preserved on the La Chapelle skeleton). Boule (1913) reconstructed the skeleton with an incompletely erect posture and concluded that the Neanderthals displayed numerous morphological specialisations that separated them, taxonomically, from Homo sapiens and indicated that they were an evolutionary "dead-end". Both Boule's reconstruction and overall conclusions can be seen as a consequence of his intellectual background (Hammond 1979, 1980, 1982).

The 1920s, 1930s and 1940s were marked by a proliferation of fossil discoveries such as the australopithecine material from South Africa, the Homo erectus material from China and Indonesia, and archaic Homo sapiens material such as that from Steinheim in Germany, Swanscombe in England and Mount Carmel in Israel (Oakley et al. 1971).

Boule's results indicated that human evolution must have involved more than a single unilinear progression, and that the "true" ancestors of modern man had not been found. Doubt about the morphological affinities of the "Pithecanthropus erectus" remains from Indonesia continued, because they too displayed primitive morphological specialisations. The expulsion of the Neanderthals from human ancestry also cast doubts on the possible ancestral status of the "Pithecanthropus erectus" remains.

These new discoveries, especially the archaic Homo sapiens or "progressive" Neanderthal remains which were easier to accept as possible humans ancestors than the "classic" Neanderthals, provided a quite different picture of human evolution from that which had been worked out from the interpretation of the Piltdown material. The australopithecine and Homo erectus remains were initially assigned to extinct branches on the evolutionary tree of modern humans because they were thought to be too "primitive" to have evolved into anatomically modem humans in what appeared at that time to be a relatively short period. However, these remains indicated that a large brain developed quite slowly, whereas features of the jaw and teeth changed over a much shorter period.

The idea that the Neanderthals formed a contemporary side branch to the direct line of human evolution became generally accepted in western Europe (e.g. Keith 1915, 1925). This general acceptance was influenced, to a large extent, by the perception that there was not enough time to transform the morphologically archaic and primitive Neanderthals into anatomically modern Europeans (Hammond 1979).

2

By the late 1940s many scholars who had previously accepted the Piltdown remains began to question their usefulness and/or validity (Weiner 1955). In 1949, Marston stated in relation to the Piltdown finds It is clear that many mistakes have been made by many highly qualified and highly placed men. To err is human and none of these men have been divine. Let the mistake be recognized. (quoted in Millar 1974:198)

Homo sapiens developed: The Presapiens, Spectrum, PreNeanderthal and Neanderthal Phase Hypotheses.

Boule's contention that the Neanderthals were an evolutionary dead-end was developed into the Presapiens Hypothesis by Vallois (1954), who identified two hominid lines in Europe. One line typified by the hominids from Swanscombe and Fontechevade, which was said to have small brow-ridges, gave rise to Homo sapiens, while the other line typified by the Steinheim and Saccopastore crania, which was said to have large brow-ridges, gave rise to the Neanderthals.

A critical re-examination of the Piltdown finds was undertaken, which found that Piltdown Man was a deliberate forgery and hoax (Weiner et al. 1953). The remains had been stained so as to resemble fossils, the cranium appeared to be of Upper Pleistocene age, whereas the mandible, canine and molars belonged to a modem orangutan, and the molars had been filed down.

Weiner and Campbell ( 1963) developed the Spectrum Hypothesis, which proposed that following the Homo erectus stage there was a spectrum of varieties. They suggested that towards the end of the Upper Pleistocene distinctive populations of Neanderthals and Rhodesian type developed, along with "intermediate" forms, such as those from Mount Carmel and Krapina, and populations of anatomically modem Homo sapiens which were the only group to survive.

The New Synthesis and the Rejection of Old Ideas During the 1930s and 1940s the modem synthesis or NeoDarwinian evolutionary theory developed when Mendelian population genetics and Darwinian natural selection were combined (Eldredge and Tattersall 1982; Gould 1982; Huxley 1942; Mayr 1980; Weiss and Chakraborty 1982). It is called a synthesis because it integrated the previously separate fields of genetics, ecology, systematics and palaeontology. This theoretical development initiated a change from the typological thinking of the first half of the century to more populational thinking (Mayr 1982). Mayr (1950), for example, reduced the plethora of hominid generic names down to two (Australopithecus and Homo) and highlighted the amount of geographical variation and sexual dimorphism present in the hominid fossil record. As Trinkaus ( 1982d) has noted, this theoretical shift is evident in palaeoanthropology with the attempt by Howell (1951, 1952) to explain the development of the unique Neanderthal morphology in populational and adaptational terms.

The Pre-Neanderthal Hypothesis as developed by Howell (1951, 1952, 1957), and elaborated upon by others such as de Lumley (1973), identified an early "generalised" group of Neanderthals in Europe that gave rise to both the later Neanderthals and anatomically modem humans. A group of these "generalised" Neanderthals became isolated in Europe due to the onset of the last glacial and developed the "classic" Neanderthal morphological configuration, while elsewhere anatomically modem Homo sapiens developed from the "generalised" Neanderthals and subsequently spread into Europe replacing the "classic" Neanderthals. The Neanderthal Phase of Man Hypothesis, which was popular with eastern European scholars, was initially based on the placement of the existing hominid record into a unilinear evolutionary sequence (Coon 1962; Hrdlicka 1927, 1930; Weidenreich 1940; 1943a, 1943b, 1947). Brace (1964) updated and re-emphasised the Neanderthal Phase of Man Hypothesis and stressed that European and western Asian Neanderthals evolved into anatomically modem humans.

The 1950s were also a period when better chronological control over the fossil record was obtained, due to the development of various radiometric dating techniques such as the radiocarbon and potassium-argon dating methods (Bishop and Miller 1972). These developments helped clarify chronological problems with hominid material such as the australopithecines, and allowed fine-tuning of the hominid evolutionary sequence.

These four hypotheses have been refined into two competing hypotheses - the African Origin or Replacement Hypothesis and the Regional Continuity or Multiregional Hypothesis. The African Origin or Replacement Hypothesis proposes that anatomically modem Homo sapiens evolved in one main area from where they radiated out and rapidly replaced the local populations in all other parts of the Old World. This explanation assumes that there was a relatively recent common Homo sapiens ancestral population. The area where anatomically modem humans originally developed is generally given as sub-Saharan Africa (e.g. Howells 1973a, 1974a, 1974b, 1976a, 1981a; Howell 1998; Stringer 1974b, , 1982, 1985, 1989a, 1989b, 1990, 1992, 1998a; Stringer and Andrews 1988; Stringer and Brauer 1994; Stringer et al. 1984). The major issue with this explanation is that it proposes a total replacement of indigenous inhabitants throughout the Old World following the migration of Homo sapiens from Africa.

The announcement of the Piltdown hoax prompted a revaluation of the ancestors of modem humans, with special attention given to the Neanderthals. At a symposium to celebrate the centenary of the recognition of the first Neanderthal remains, Straus and Cave (1957) criticised and tried to correct the erroneous image of the Neanderthals created by Boule (1913). They stated, if a Neanderthal ...could be reincarnated and placed in a New York subway-provided that he were bathed, shaved and dressed in modem clothing-it is doubtful whether he would attract any more attention than some of the other denizens. (1957:359) From this re-evaluation of the course of human evolution four major theories for the origin of anatomically modem

The Regional Continuity or Multiregional explanation emphasises regional morphological continuity and argues 3

that anatomically modem humans developed from groups that had already differentiated from each other (Homo erectus-archaic Homo sapiens). Weidenreich (1940; 1943a, 1943b, 1947) developed this explanation by formulating a polycentric interpretation of hominid evolution based on vertical and horizontal differentiation of the material. He proposed four separate (although not necessarily isolated) origins of anatomically modem humans that correspond to the major modem geographical divisions (Eurasian, Mongolian, African and Australian). Allowance was given (due to the dating of the hominid sample available to W eidenreich) for the acceleration or retardation of the evolutionary development in the four lines. Weidenreich (1947:201) developed a diagram on which he identified evolutionary phases through which hominids passed to become anatomically modern. The diagram did not present a simple unilineal parallel development in the four evolutionary lines in that Weidenreich represented not only vertical lines, but also horizontal and diagonal ones. Howells (1959) simplified and diluted Weidenreich's explanation by referring to it as the "candelabra theory".

Not only did the number of hominids increase in the 1980s and 1990s, but also the number of taxonomic units into which they were placed . Hominid phylogeny changed from a tree to a multi-branched bush (White 2003). Like the 1950s, the late 1980s and 1990s was also a period when better chronological control over the fossil record was obtained, due to the application of new or refined radiometric dating techniques to the archaeological record including uranium-series, electron spin resonance, accelerator mass spectrometry radiocarbon dating and thermoluminescence. These developments clarified the chronological placements of many hominids, especially those in the Levant from the sites of Tabiin, Skhiil and Jebel Qafzeh. It is also worth noting that there have been refinements in the calibration of radiocarbon ages that have generally underrepresented "true ages" (e.g. Bard et al. 1990). This period has also seen the interpretation of analyses of genetic data has entered the debate on the origin of anatomically modem humans. Mitochondrial DNA (mtDNA), especially, has been used to study human evolution because it accumulates mutations faster than nuclear DNA and is inherited maternally and so can be viewed as a genealogy linking maternal lineages in modem human populations to a common ancestral female. Mitochondrial DNA has also been recovered from Neanderthal remains, anatomically modern Upper Palaeolithic skeletons and ancient Australians. The mtDNA sequence results have been interpreted as indicating genetic continuity between anatomically modem Upper Palaeolithic and modem populations and a discontinuity with the Neanderthals who did not contribute to the mtDNA pool of modem humans, whereas one ancient Australian, Lake Mungo 3, was separated from the other Australian and modem samples and so may have diverged before the most recent common ancestor of contemporary human mtDNA sequences. These mtDNA results have been criticised on the grounds of methodology, interpretation and whether noncontaminated mtDNA can actually be sequenced from prehistoric hominid material. Genetic analyses hold great prospect for adding substantially to the population history of anatomically modem humans in the future.

Wolpoff et al. (1984) refined Weidenreich's polycentric interpretation of hominid evolution and developed the Multiregional Hypothesis. The Multiregional Hypothesis contends that regional variations have developed at the species' peripheries (via Thome's [1981a] centre and edge hypothesis) and are maintained by a balance between gene flow and selection, which caused the formation of morphological clines (Thome and Wolpoff 1991b; Wolpoff et al. 1984; Wolpoff et al. 2000). These clines could be local, regional, and/or across the entire species range. There were not isolated morphological clades because inherent within this explanation is gene flow across the species' distribution. It is the balance between selection, gene flow and genetic drift that underpins the Multiregional Hypothesis (Wolpoff 1985, 1989b; Wolpoff et al. 1984) and which, it is argued, has resulted in a long-lasting dynamic system of morphological clines that characterise the multiregional distribution of the polytypic hominid lineage. The Current Phase The 1970s were marked by the discovery of many PlioPleistocene hominids from Africa (Day 1986). These discoveries focussed scientific attention on the early stages of hominid evolution and drew attention away from Europe and the appearance of modem humans.

Re-emphasis on the problem of the origin of anatomically Homo sapiens, refined dating schemes and the addition of genetic analyses has not resulted in the resolution of which of the Multiregional Hypothesis or the Replacement Hypothesis best explains modem human origins. Willermet and Clark observed It has become evident that the two camps are operating with different underlying preconceptions about what causes pattern in the human biological record, and how pattern should be measured. (1995:487) and that the two hypotheses each represent distinct paradigms. Therefore, unlike in the late nineteenth and earlier twentieth centuries where the interpretations of the hominid fossil record was greatly influenced by the intellectual predilections of the society at that time, it is now more the preconceptions, biases and preferred paradigm of particular scholars that is framing the interpretation of the origin of modem humans (cf. Frayer et al. 1993, 1994; Stringer and Brauer 1994). Willermet and Clark (1995 :487)

During the 1980s the evolutionary spotlight was redirected on to the -problem of the origin of Homo sapiens, as is clearly demonstrated by the number of symposia, articles and books that have dealt with this topic (e.g. Aiello 1993; Akazawa et al. 1998; Brauer and Smith 1992; Gowlett 1987; Mellars and Stringer 1989; Ronen 1982; Smith and Spencer 1984; Trinkaus 1983, 1989). Additional hominid fossils were recovered during this period that have direct relevance to modem human origins. Hominid material that had been overlook at Feldhofer Neanderthal site were re-excavated (Schmitz et al. 2002), while previously excavated hominids such as Le Moustier 2 and Combe Capelle, which had be "lost" were rediscovered (Hoffmann and Wegner 2002; Maureille 2002).

4

Gingerich ( 1984) suggested that stasis can be viewed as gradualism at zero rate. He also argued that abse~ce of evidence (gaps in the fossil record) is absence of evidence, and not, as the punctuated equilibria model implies, evidence of absence.

even suggested that modem human origins research was in "a classic case of paradigm crisis".

Tempo and Mode - Phyletic Gradualism vs. Punctuated Equilibria

Behrensmeyer (1976) and Cronin et al. (1981) have also documented the very small percentage of the total population of hominids that have actually been preserved at Koobi Fora and Omo Shungura, respectively. This demonstration of the inadequacies of the hominid fossil record casts doubt on the contention of the punctuated equilibria model that gaps in the fossil record are real.

The mode of evolution has also been much debated. The Neo-Darwinian synthesis, and especially phyletic gradualism, which argued that species arise by the transformation of the entire ancestral population (or a large part of it) into its modified descendants, has been challenged (for example Eldredge and Gould 1972; Eldredge and Tattersall 1982; Gould 1980a, 1980b, 1982, 1983, 1985; Gould and Eldredge 1977, 1983, 1986; Stanley 1978, 1979, 1982, 1985; Vrba 1980, 1984).

Gould and Eldredge have also argued that ...there is no direct evidence for gradualism within any hominid taxon ...Each species disappears looking much as it did at its origin ... (1977: 135) This view has been both supported (Delson 1981, 1985b; Delson et al. 1977; Eldredge and Tattersall 1982; Pilbeam 1975, 1986; Stanley 1979, 1981, 1985; Vrba 1984), and contested (Bilsborough and Wood 1986; Brace 1981; Cronin et al. 1981; Stringer 1984b; Tobias 1983; Wolpoff 1984).

Eldredge and Gould ( 1972) argued that within-species (phyletic) evolution was too slow to produce the major evolutionary events such as speciation. They felt that evolution was characterised by rapid speciation events (punctuation), separated by long periods of morphological stasis during which species change only slightly, if at all, and the change that does occur consists of non-directional fluctuations in form (equilibrium). Most of the evolutionary change in punctuated equilibria, as this model has been called (Eldredge and Gould 1972; Gould and Eldredge 1977), is concentrated in rapid, often geologically instantaneous, speciation events that occur in small isolated peripheral populations. Exponents of punctuated equilibria also contend that breaks in the fossil record are "real data". Their explanation separates macroevolution (large scale evolutionary change such as speciation) from microevolution ( small scale change within existing species), whereas phyletic gradualism closely links the two. The African Orig~ or Replacement Hypothesis has been portrayed as a rapid speciation event that occur in small isolated populations in Africa.

Cronin et al. ( 1981) addressed the evidence put forward by Gould and Eldredge ( 1977) in support of the contention that the hominid fossil record upholds the punctuated equilibria model and concluded that a hypothesis of phyletic gradualism was more parsimonious. Cronin et al. ( 1981) plotted mean body weight and cranial capacity through time and calculated regression lines which indicated that there was no evidence of marked stasis or punctuation events. However, Turner (1985) has criticised the arguments put forward by Cronin et al. (1981), while Stanley (1985) has discussed the problems of using body size in this type of analysis. Homo erectus has become a focal point for the debate over tempo and mode in hominid evolution. Different studies of this hominid species have found evidence of gradual change (Allen 1982; Bilsborough 1976; Bilsborough and Wood 1986; Bridges 1984; Cronin et al. 1981; Leigh 1988; Lestrel 1975; Lestrel and Read 1973; Stringer 1984b; Tobias 1971; Wolpoff 1984, 1986b), while others have found evidence of stasis (Day 1982, 1984; Delson 1981, 1985b; Howells 1980; Kennedy 1983; Rightmire 1981a, 1982; Walker 1984).

The punctuated equilibria model has not been accepted by all, and a gradualist-punctuationalist debate has developed. For example, it has been argued that punctuated equilibria are consistent with the Neo-Darwinian synthesis and Wrightian genetic microevolutionary models, and that it had been predicted by Darwin (Charlesworth et al. 1982; Dawkins 1985; Gingerich 1984; Lande 1985; Lewin 1986; Lovtrup 1981; Newman et al. 1985; Rhoads 1983; Tobias 1983).

Cronin et al. ( 1981) argued that the early African Homo erectus crania, such as KNM-ER 3733, differed from later examples from Africa, Indonesia and China, and so the available material did not support the contention that Homo erectus displayed morphological stasis.

The indiyidual elements of the punctuated equilibria model, such as the identification of periods of stasis and punctuation, have also been questioned. The geologically instantaneous punctuation events may involve 50,000 years, which on a microevolutionary scale is a long time, during which marked morphological changes could occur (Stebbins and Ayala 1981; Williams 1987). Williams (1987) argued that the fossil record generally underestimates the rates of evolution that have occurred and that within-species evolution of living species is sufficient to account for punctuational events in the fossil record. Gingerich (1983, 1984) argued that if punctuation and stasis were properly scaled, they would be almost the same because when high rates of change calculated over short intervals and low rates of change calculated over long periods are corrected for differences in measurement intervals, they would be essentially the same (but see Gould 1984 and Stanley 1985).

Rightmire ( 1981a) plotted the mean cranial capacity, biauricular breadth, breadth of the first lower molar and mandibular robusticity of chronologically grouped individuals against time, and calculated the least squares linear regression. He found that, although the four variables fluctuated throughout the existence of Homo erectus, the rate of change was not significant. Rightmire (1981a) concluded that Homo erectus exhibited little morphological change during its existence. Levington ( 1982) challenged Rightmire's (1981a) approach and posed the question that, given the inadequacies of the data set, could one expect to 5

see any evolutionary trends? Rightmire (1982), although not abandoning his overall conclusion, conceded that there were grounds for doubting the results of his analysis. Bilsborough and Wood (1986) have argued that Rightmire's (1981a) conclusions were sweeping, considering be only used four variables, and that there were problems with the variables used. Allen (1982) reanalysed Rigbtmire's (1981a) data, but plotted individual hominids instead of grouped means (the exact dating of the hominids is therefore a problem) and identified significant evolutionary change within Homo erectus. This suggests that Rightmire's (1981a) method (plotting group means) may have influenced bis results. Rightmire's (198 la) results are also interesting when one considers that he included in bis analyses hominids that are not generally classified as Homo erectus, for example, the Petralona cranium is generally classified as archaic Homo sapiens (Chapter 5; Habgood 1988, 1989e; Stringer 1984a; Stringer et al. 1979). If the punctuated equilibria model was applicable to hominid evolution, one would have expected a major punctuation event in the sequence between the Homo erectus material and the Petralona cranium, which Rightmire (1981a) did not find. In a later analysis of cranial capacity alone, Rightmire ( 1985) plotted individual crania, and reaffirmed his earlier conclusion that there was little evidence for rapid morphological change within Homo erectus. From this analysis, however, Rightmire (1985) suggested that there was an increase in the rate of change in cranial capacity between Homo erectus and archaic Homo sapiens. Alternately, Lestrel (1975; Lestrel and Read 1973) and Stringer ( 1984b), Leigh ( 1988) and Clausen ( 1989) have identified more definite trends in cranial capacity increase within Homo erectus.

differences, and found that the profiles for early African Homo erectus specimens fell within the shape envelope (range of variation of the profiles) for later Homo erectus (excluding the Ngandong sample). Stanley (1985) used Walker's (1984) study to support the punctuated equilibria model for hominid evolution. Walker's {1984) analysis, however, has shortcomings in that it would appear to be a relatively inefficient way to investigate changes in cranial shape. Walker (1984) eliminated size differences and so removed the effects of endocranial enlargement, which is a significant phenomenon in hominid evolution. Also, although on Walker's (1984:Fig. 4) diagram the standardised coronal and sagittal profiles of early (Koobi Fora) Homo erectus fell within the standardised profile envelopes for late Homo erectus, there appears to be definite variation of the profiles evident. Cronin et al. ( 19 81) have also demonstrated that the KNM-ER 3733 cranium differs in many features from later Homo erectus specimens. Walker (1984) also concluded that archaic Homo sapiens were a larger version of this ancestor, with little shape change, but if punctuated equilibria had occurred in hominid evolution, one would have expected major shape changes between archaic Homo sapiens and Homo erectus. Anton (2002) found differences on a "shared morphological bauplan" between the early Indonesian hominids and later Chinese and Indonesian Homo erectus material. As outlined above, Gould and Eldredge ( 1977) argued that hominid species are distinct entities with definite beginnings and ends, and that they disappear looking much the same as they looked at their beginning. Although dating issues remain, this contention does not seem to be applicable to Homo erectus. Early examples of Homo erectus such as KNM-ER 3733 and KNM-ER 3883 are morphologically different from later examples. These two crania are much smaller than Olduvai H9 and differ in most metrical characters (Leakey and Walker 1985; Rightmire 1979, 1984b) and many morphological features. There is significant morphological variation among Homo erectus material from Africa, China and Indonesia, which has prompted some scholars to suggest that they represent difference species {Andrews 1984a, 1984b; Groves 1989; Stringer 1984a; Wood 1984).

Wolpoff ( 1984, 1986b) conducted two analyses to examine whether Homo erectus provides evidence of stasis or gradual change. In his initial study, W olpoff ( 1984) analysed thirteen cranial, dental, and mandibular variables for a narrowly defined Homo erectus sample (which excluded the Ngandong specimens), divided into three temporal subsets ( early, middle and late). He found significant evolutionary differences between the early and late samples, with the cranial vault expanding while the masticatory structure was reduced. He also found that change between a late Homo erectus group and an early archaic Homo sapiens sample was in the same direction as the change within Homo erectus. Delson (1985b:762-3), however, has stated that although Wolpoffs results "are superficially persuasive ...there are too many potential problems for bis analysis to be convincing". The problems in Wolpoff s ( 1984) analysis that Delson ( 1985b) · cited included the questionable identification of some of the individual specimens, the effects of sexual dimorphism and the use of time bands instead of age estimates, but these problems are also present in Rightmire's (1981a) initial study. Wolpoff (1984) also contends that the rate of change that he found within Homo erectus was at or above the fossil vertebrate average. Wolpoff ( 1986b) later undertook another analysis leaving out some individuals and adding others into three temporal groups, while reducing the number of variables to four. Wolpoff (1986b) found that the patterns and magnitude of change evident in this new analysis closely resembled his earlier results (Wolpoff 1984). Walker (1984) has also suggested that Homo erectus evolved quickly and remained relatively unchanged, except for size changes, for over a ·million years. Walker ( 1984) plotted sagittal and coronal profiles standardised to remove size

Andrews (1984a, 1984b), Groves (1989), Stringer (1984a) and Wood ( 1984) cladistically separated the African and Asian Homo erectus specimens, arguing that the former have more claim to the ancestry of anatomically modem humans than the latter. Wood (1984; Bilsborough and Wood 1986) even separated KNM-ER 3733 and KNM-ER 3883 from Olduvai H9, grouping the latter with Asian Homo erectus. Stringer (1984a) also found some individuals within the Ngandong sample differ from other Homo erectus specimens in the direction of anatomically modem humans. Bilsborough (1976) also suggested that the Zhoukoudian Homo erectus remains should be included in Homo sapiens, and so differentiated from other Homo erectus material. Also, where multiple individuals have been recovered from the same site, as at Dmanisi and Zhoukoudian, there is generally both similarities on a common morphological theme as well as individual morphological variation. These results indicate that there are marked differences within the Homo erectus species and so there is no real evidence for morphological stasis. Rather, there is clear evidence of variation and change. 6

The hominid fossil record from Homo erectus through to anatomically modern humans does not provide any clear evidence of stasis, but rather documents gradual morphological change. For this reason the latter stage of human evolution (archaic Homo sapiens through to anatomically modern Homo sapiens) has been viewed from the perspective of phyletic gradualism. As Delson, a supporter of punctuated equilibria (Delson 1981, 1983), has stated with respect to punctuated equilibria and gradualism Such a discrete dichotomy is surely unrealistic as the question is more one of relative frequency than of near-absolute dominance of one or other mode of evolution, especially given the problems involved in assigning many cases studied to a particular mode. (1983:101)

The late Middle Pleistocene hominid record from Europe documents the gradual development of the Neanderthal morphological configuration over some 200,000 years (Chapter 5; Ruhlin 1984; Smith 1985; Stringer 1985; Wolpoff 1980a, 1980b, 1989a), and not a punctuation event. Evolutionary trends have also been identified within the Neanderthals (Brace 1979; Frayer 1978; Smith 1982, 1984; Smith and Paquette 1989; Smith and Ranyard 1980; Wolpoff 1980a) and early anatomically modem humans (Frayer 1977, 1978, 1980, 1981, 1984; Jacob 1985). Does the transition to Homo sapiens provide evidence of a punctuation event? If phyletic gradualism typifies the transition from Homo erectus to Homo sapiens, late members of the former species should be more similar to Homo sapiens than to earlier Homo erectus specimens, while the early archaic Homo sapiens specimens should display a mosaic of Homo erectus and Homo sapiens features. The boundary between Homo erectus and Homo sapiens is not easy to delineate morphologically, as Gould and Eldredge ( 1977) would have us believe it should be. As mentioned above, Bilsborough (1976) suggested placing the Zhoukoudian Lower Cave material within Homo sapiens, while Stringer (1984a) has found that although the Ngandong sample should be regarded as Homo erectus some individuals display features more typical of Homo sapiens. Brauer and Mbua 1992; Kennedy 1991; Rightmire (1998), The difficulty in identifying this boundary has prompted some authors to suggest that Homo erectus should be included within the species Homo sapiens (e.g. Bonde 1981; Hemmer 1969; Jelinek 1978, 1980a, 1981; Wolpoff et al. 1984).

Morphological Relationships - Cladistics vs. Phenetics There are two major techniques for estimating relationships between hominid taxa when using morphological data cladistic and phenetic analyses. Many supporters of punctuated equilibria have adopted cladistics or phylogenetic systematics, as developed by Hennig ( 1966). What follows is not meant to be a detailed explanation or critique of cladistic analysis (see Wiley 1981), which has a sound logical basis, but rather is a discussion of some of the limitations of a pure cladistic approach when applied at low taxonomic levels and in evaluating the latter stages of the evolution of the genus Homo. Cladistic analysis is the analysis of branching ( cladogenetic or cladogenesis: pertaining to the branching sequence in evolution [Ashlock 1974; Holmes 1980]). The cladistic method has required more explicit definitions of morphological traits. Cladistics identifies "different kinds of similarity" (Eldredge and Tattersall 1975), namely: pnnnt1ve, ancestral or "plesiomorphic" traits which are inherited without modification from a remote ancestor; and derived, advanced or "apomorphic" traits which are transformations of the primitive condition (Delson et al. 1977; Hennig 1966; Skelton et al. 1986; Trinkaus 1990). It is argued that shared primitive character states, _or "symplesiomorphies", do not contain specific information about relationships and that the only useful traits for reconstructing relationships are shared derived character states or "synapomorphies" (e.g. Andrews 1984a, 1984b, 1986; Bilsborough and Wood 1986; Delson 1978; Delson et al. 1977; Eldredge and Cracraft 1980; Eldredge and Tattersall 1975, 1982; Foley 1989; Lieberman 1995; Hennig 1966; Skelton et al. 1986; Tattersall and Eldredge 1977; Wood 1984; Wood and Chamberlain 1986). A trait is "derived" in the first species or group to have it, but then becomes a primitive character for the descendants of that species or group. The traits that are identified as "derived" are weighted at the expense of those character states that have been identified as primitive.

If phyletic gradualism did occur, there should also exist hominids around the world that display "transitional" erectuslsapiens features. The hominids from Ngandong, Saldanha, Bodo, V ertesszollos, Bilzingsleben, Arago, Ndutu, Narmada, Kabwe 1, Omo 2, and Petralona crania are without doubt difficult to classify (Chapters 2, 4, 5 & 6). The difficulty of their attribution has been highlighted in multivariate analyses (Habgood 1982, 1984b, 1988; Habgood and Walker 1986) where some of the Ngandong specimens, which are most often classified as Homo erectus, grouped with the Petralona crania, which is most often classified as archaic Homo sapiens. This material is clearly different from those early Homo erectus crania such as KNM-ER 3733 and KNM-ER 3883, and from later Homo sapiens (including Neanderthals and anatomically modern humans). Gould and Eldredge have argued that Small numbers and rapid evolution virtually preclude the preservation of speciation events in the fossil record ... (1977: 117)

Yet these early Middle Pleistocene hominids seem to preserve the legacy of a speciation event or events. One would also not expect to find "transitional" specimens throughout the geographic distribution of Homo erectus, as have been found, if speciation occurred in a small peripheral population as the punctuated equilibria model argues (Eldredge and Gould 1972; Gould and Eldredge 1977). What the material would seem to indicate is a gradual morphological transition or transitions from Homo erectus to Homo sapiens.

As Tattersall and Eldredge (1977) have outlined, after a morphocline has been established and the direction of change determined, a cladogram can be produced. This is a branching diagram that depicts the pattern of distribution of derived character states within a group of organisms (usually a monophyletic group [Ashlock 1974; Holmes 1980]). It is assumed that the sharing of derived character states indicates 7

group) into two or more daughter species (sister species of one another) and assumes that traits change in consistent ways through time and once lost cannot reappear as evolutionary reversals (Brace 1981; Habgood 1989c; Trinkaus 1990, 1992; van Vark 1978). Trinkaus (1990) pointed out that morphological similarities due to homoplasies (including convergences, parallelisms and reversals) are often not considered in cladistic analyses and that it is inherently difficult to distinguish homoplasies from shared ancestral traits ( symplesiomorphies) or shared derived traits ( synapomorphies).

an evolutionary relationship, although this is unspecified on the cladogram in that no ancestor-descendant relationships are postulated because chronological data are not considered (Delson et al. 1977; Eldredge and Tattersall 1975; Tattersall and Eldredge 1977; Wiley 1979). Whilst the cladistic approach has the advantaged of requiring explicit definition and assessment of morphological traits, it also has methodological limitations (e.g. Ashlock 1971, 1972, 1974; Brace 1981; Charig 1981; Darlington 1970; Gingerich 1979; Habgood 1989c; Halstead 1978; Halstead et al. 1979; Harper and Platnick 1978; Mayr 1965a, 1965b, 1969; Panchen 1979; Szalay 1977; Trinkaus 1990, 1992; van Valen 1978).

Cladistic classifications can also be inherently unstable because the reclassification of a single character state from an inferred synapomorphy to an inferred symplesiomorphy, or vice versa, or the discovery of a new fossil, can necessitate major changes to a cladogram.

Cladistic analyses generally use only a few synapomorphies traits that have been heavily weighted by the cladist. Therefore, much information, in the form of the features inferred to be symplesiomorphies (which are not considered) , can be lost. Also, as Eldredge and Tattersall have stated Usually, however , it is found that some taxa appear to be allied with certain others in one or more characters, but with yet others when different characters are considered . (1975:230)

For these reasons cladistic classifications often fail to mirror evolutionary descent (Ashlock 1974; Brace 1981; Panchen 1979; Szalay 1977). Underlying many cladistic classifications is the assumption that if at least one character state that is inferred to be derived is shared by two groups, then they must have had a more recent common ancestor and so be more closely related to each other than to other groups ( e.g. Cracraft 1981; Delson 1978; Delson et al. 1977; Eldredge and Cracraft 1980; Eldredge and Tattersall 1975, 1982; Skelton et al. 1986; Tattersall and Eldredge 1975; Wood 1984; Wood and Chamberlain 1986). This assumption is also inherent in cladistic classifications based on individual character analysis (e.g. Wood 1984). Most cladistic analyses also do not consider populational variability, although Stringer (1984a; Day and Stringer 1982) has suggested that a majority of (or at least 50% of) a range of characters needs to be present. This would seem a more reasoned approach in that it does consider morphological variation.

Cladistic analyses also have difficulties with time, which, as mentioned above, is not represented on cladograms. Some cladists (e.g. Delson et al. 1977; Vrba 1980) argue that ancestor-descendant relationships cannot be deduced from cladograms (but see Wiley 1979). However, it is the investigation of time and ancestor-descendant relationships that evolutionary studies are interested in and so one could be excused for thinking that cladograms have only limited use in such studies. Foley ( 1987) even added stone tool type to a cladogram of hominid evolution and suggested they the manufacture of stone tools could be regarded as a unique apomorphic trait for the genus Homo .

What follows are some examples of the problems encountered when a strict cladistic approach based on individual character analysis that does not consider morphological variation is adopted.

Tattersall and Eldredge (1977:207) suggested that phylogenetic trees should always be based on cladograms, although they were "unable to provide a clear-cut methodology for tree formation". There is also the limitation that an enormous number of phylogenetic trees can account for each cladogram, with Harper and Platnick ( 1978) proposing that the number was actually infinite (Eldredge and Tattersall 1975; Felsenstein 1978; Harper and Platnick 1978; Tattersall and Eldredge 1977). This limits the usefulness of cladograms for evolutionary studies. Phylogenetic trees also contain more information than cladograms because they express both ancestor-descendant (anagenetic) and branching-sister group (cladogenetic) relationships. They therefore permit a greater insight into evolutionary relationships (Szalay 1977). Rightmire (1996:824) suggested that under cladistics, as "a species is diagnosed more completely, it will be harder to place it in any ancestor-descendant sequence".

The Neanderthals Stringer et al. ( 1984:Table 2) have listed features that they proposed as "autapomorphies" and common characteristics for the Neanderthals. Not all Neanderthals , however, display all of these features.

The horizontal-oval form of mandibular foramen is found on a large proportion of Neanderthal mandibular rami: • 46.2% of European and western Asian Neanderthals (Smith 1978b); • 61.5% of adult European Neanderthals (Trinkaus 1983a); • 56.3% of early European Neanderthals (Stefan and Trinkaus 1998); • 41.7% of later European Neanderthals (Stefan and Trinkaus 1998); • 52.6% of European Neanderthals (Frayer 1992); and • 16.7% of western Asian Neanderthals (Stefan and Trinkaus 1998).

A major limitation of the cladistic approach is that it has difficulty in accounting for mosaic evolution (differing rates and times of change for different traits within a single line or between related lines), continuing change after a given branch point (evolutionary parallelism and convergence) and phyletic evolution (anagenesis), because cladistics characterises phylogeny as the splitting of one species (or 8

In cladisitic terms the horizontal-oval form of mandibular foramen is a suggested European shared-derived character as it is also found on European Upper Palaeolithic anatomically modern Homo sapiens mandibular rami: • 23% of European Upper Palaeolithic anatomically modern humans (Smith 1978b); • 26.7% of early European Upper Palaeolithic anatomically modem humans (Stefan and Trinkaus 1998); • 44.4% of early European Upper Palaeolithic anatomically modem humans (Frayer 1992); • 5.3% of late European Upper Palaeolithic anatomically modem humans (Frayer 1992); and • 1.9% of European Mesolithic material (Frayer 1992). It is worth noting that the Tabtin C2 and Kebara 2 mandibles are the only western Asian archaic and early modern individuals, including western Asian Neanderthals and the Skhiil and Jebel Qafzeh material, that have the horizontaloval mandibular foramen form (Stefan and Trinkaus 1998). The horizontal-oval mandibular foramen pattern may also be present on the Homo erectus mandible Olduvai H22 and Zhoukoudian H-1, although it is not present on the remainder of the Zhoukoudian mandibles, the Ternifine 2 and 3 mandibles or the early European Mauer mandible (Groves 1989; Smith 1978b).

Aborigines. Some of the late Pleistocene Coobool Creek crania also have this foramen, although the Lake Mungo 1 cranium does not (Pardoe 1988b). The Ngandong calvaria also lack an inferior petrosal sinus (Pardoe 1988b). Pardoe ( 1988b) did not find this feature described on crania from outside Australia, Melanesia and the Pacific region. In a cladistic analysis, prehistoric Australian Aboriginal crania would have a unique derived "autapomorphic" trait, ~hich would separate them from other modem human groups that lack an inferior petrosal sinus. However, the Australian Aborigines are clearly anatomically modem Homo sapiens (Brown 1990; Wolpoff 1986a) and cannot be taxonomically separated from other anatomically modem humans. Asian Homo erectus As noted above, some authors utilising a cladistic approach have concluded that Asian Homo erectus sensu stricto specimens show a concentration of derived characters that indicate they contributed little, if anything, to the ancestry of anatomically modem Homo sapiens (cf. Andrews 1984a, 1984b; Bilsborough and Wood 1986; Stringer 1984a; Wood 1984; see also Delson et al. 1977; Eldredge and Tattersall 1975, 1982; Groves 1989; Tattersall and Eldredge 1977; but see Habgood 1989c; Trinkaus 1990, 1992).

Wood (1984; Bilsborough and Wood 1986) concluded that KNM-ER 3733 and KNM-ER 3883 did not belong to Homo erectus because they lacked the autapomorphic features of Homo erectus, but he did include the Olduvai H9 calvaria from Africa along with Asian material in Homo erectus sensu stricto. This conclusion would appear to be based, predominantly, on the greater robusticity of Olduvai H9 when compared to the two other crania. KNM-WT 15000 could be grouped with Olduvai H9 in that Brown et al. suggested It is conceivable that this individual, had it lived to maturity, might have developed as strong and robust a cranium as that of Olduvai Hominid 9. (1985:789) If this taxonomic attribution of the hominids was correct there would have been two contemporary species, one robus~ and classified as Homo erectus and the other more gracile and classified as non-Homo erectus (Homo ergaster), living around Lake Turkana. What is more probable is that there was a single sexually dimorphic and morphologically variable species, Homo erectus, inhabiting the area.

Some degree of taurodontism (enlargement of the pulp cavities of the molars) was found on 90.4% oflower molars from Krapina (Kallay 1963, 1970b) and is quite common for other Neanderthals (Stringer et al. 1984; Trinkaus 1983a), while also being found among modem Homo sapiens (Blumberg et al. 1971; Coon 1962). These two character states cannot, therefore, be regarded as synapomorphies for the Neanderthals (sensu lato) as they are not found on every member of this group and are also found on members of another group, anatomically modern Homo sapiens (see also Lieberman 1995). Also, mid-facial prognathism and a suprainiac fossa cannot be used as synapomorphies for the Neanderthals (sensu lato), as they are present on earlier hominids outside Europe Kabwe 1 displays the former character state and Eliye Springs 11693 the latter - and so in a strict cladistic sense they also must be regarded as symplesiomorphies. However, it could be argued that these two features are synapomorphies and that Kabwe and Eliye Springs 11693 are more closely related to the Neanderthals than to other African · hominids that lack these traits. More detailed morphological comparisons indicate this to be improbable (Chapter 2; Habgood 1988).

Stringer (1984a:131) also found KNM-ER 3733 and KNMER 3883 to "predominantly share plesiomorphies with Asian H. erectus, rather than autapomorphies", but observed that the assignment of KNM-ER 3733 and KNM-ER 3883 to a grade called Homo erectus was justified on phenetic grounds "but the species so constituted cannot be satisfactorily defined cladistically".

These four traits therefore, should not be used in cladistic classifications involving the Neanderthals, yet they are useful features for characterising this group (sensu lato and sensu stricto).

Andrews synthesised the papers of Wood (1984) and Stringer ( 1984a) and stated ...my provisional interpretation of the evidence would be that the African skulls formerly attributed to erectus would have been close to the line leading to sapiens and that the Asian erectus was some way removed from this lineage. ( 1984a: 172)

Australian Aborigines Another example of the problems encountered when a strict cladistic approach (cf. Wood 1984) is used, is illustrated by the Australian Aborigines. The distribution of the inferior petrosal sinus would appear to be restricted to Australia, Melanesia and the Pacific (Pardoe 1988b). Pardoe ( 1988b) found this trait in 36% of a sample of southern Victorian 9

In the same volume that contained the papers by Andrews (1984a), Stringer (1984a) and Wood (1984), Rightmire ( l 984b:83) found that the evidence suggesting overall similarity of form between Asian and African Homo erectus material was "much more striking" than the differences and that "Homo erectus seems to have been a 'real' species". Similar conclusions have been reached by other morphological studies of Homo erectus sensu lato. Brauer and Mbua ( 1992) found that a continuous range of morphological variation was evident within African and Asian Homo erectus material for the features suggested to be uniquely derived characters. Rightmire ( 1998) found that, although there was variation, facial morphology did not separate African and Asian Homo erectus material and that all of the hominids could be placed within Homo erectus. Kennedy ( 1991) concluded that, whilst it should be retained, Homo erectus was not a valid taxon because no autapomorphic features had been identified. Hublin ( 1986) has proposed that most of the derived features of Homo erectus that were identified by Andrews (1984a), Stringer (1984a), and Wood (1984), were related to an increase in the compacted bony mass of the vault and that this phenomenon was especially developed in the Asian material. There would, then, only be one autapomorphic character present on Asian Homo erectus crania, especially developed bony hypertrophy, which separated them from the African material and not four as proposed by Andrews (1984a), six as Wood (1984) concluded, or twelve as Stringer (1984a) suggested. As Skelton et al. observed It is generally not advisable to consider traits that appear to be evolving in tandem as separate, since this tends to inflate the importance of what may actually be only one phenomenon. (1986:22) As noted earlier, inherent in cladistic classifications is the assumption that if two or more individuals, groups or species share one or more derived traits relative to other species, then they must have shared a more recent common ancestor and must be more closely related to each other than either is to any other species. For example, Wood contends The logic of the cladistic system is that if two crania show any evidence of H. erectus autapomorphies, no matter how few these may be, then they must belong to H. erectus, even if they lack the characteristic combination of primitive, or shared-derived, traits. (1984:107) Andrews (1984a), Stringer (1984a), and Wood (1984) all indicated that the presence of an angular torus was a Homo erectus sensu stricto (Asian) autapomorphic character, yet its occurrence on the Ngandong crania is variable (Anton 1999; Santa Luca 1980), OH 12 from Africa appears to have an angular torus (Kennedy 1991; Rightmire 1986) and an angular torus is also found on archaic Homo sapiens crania from Africa, Europe and Asia including Bodo VP-1/1, Herto 1, Dali, Narmada, Ceprano, Atapuerca SH At Cr-4 (AT-600), Arago 4 7, Reilingen and possibly Castel di Guido 5 (Chapters 2, 5 & 7; Grimaud 1982; Kennedy 1991; Mallegni et al. 1983). WLH 50 from Australia also has angular tori. Weidenreich (1943a) described Kabwe 1 as having an angular torus , whereas Santa Luca ( 1980: 105) observed that

the elevation between the lambdoid suture and the parietal incisure "is part of the upper lateral arm of the occipital torus, which connects the former with the mastoid crest". The elevation on the posteroinferior parietals of Kabwe 1 does not morphologically match the angular tori found on Asian Homo erectus material, but the elevations can still be referred to as angular tori (see also Stringer (1984a). Weidenreich (1943b:37) stated in relation to the angular torus "The same arrangement as in Sinanthropus may occasionally be found in modem man". Lamach and Macintosh (1966, 1970), using Weidenreich's (1943a) definition (which is broader than that of Santa Luca [1980]), found angular tori on 39.4% of a male and 27.1 % of a female sample from coastal N.S.W. (34.7% of the total sample), and on 52.1 % of a male and 22.2% of a female sample from Queensland (39.1% of the total sample). Andrews (1984a) and Stringer (1984a) also list other so-called autapomorphies of Asian Homo erectus crania that are present on the material from Arago, Bodo and/or the Petralona cranium. Adopting a strict cladistic approach would mean that all of the above mentioned archaic Homo sapiens crania, the WLH 50 cranium and many modem Australian Aboriginal crania must be regarded as Homo erectus because they have at least one Homo erectus autapomorphic character (an angular torus). This material also displays many Homo sapiens autapomorphies and so must belong to that taxon as well. As noted by Stringer (1984a:139), some individuals in the Ngandong sample "certainly differ from the typical H. erectus s.s. morphology in ways approximating that of H. sapiens s.I". If one were to adopt a strict cladistic approach, the Ngandong material must belong to Homo sapiens, whereas most of the cladistic evidence suggests that they are Homo erectus (Santa Luca 1980; Stringer 1984a). Absence of a styloid process Another example of the problems that such a narrow view of morphological variation can cause is the absence of a styloid process. This feature is evident on KNM-ER 3733, KNM-ER 3883, Olduvai H9, Ndutu, Kabwe 1 Ileret ER-3884 and Atapuerca GD (Chapters 2 & 5; Clarke 1984; Leakey and Walker 1985; Rightmire 1980, 1983), but is absent on australopithecine crania (Howell 1978), the Zhoukoudian and Indonesian Homo erectus crania (Jacob 1966; Weidenreich 1943b, 1951) and east Asian archaic Homo sapiens crania from Dali (Pope 1992). ls this another feature that demonstrates that African crania are the ancestors of anatomically modem Homo sapiens crania which normally have a styloid process? A styloid process is also present on the Narmada calvaria from India ( de Lumley, M-A. and Sonakia 1985). Does the presence of a styloid process align the Narmada hominid with African Homo erectus when many other morphological features , such as the presence of an angular torus, can be paralleled on Asian Homo erectus material? A styloid process is absent or rudimentary on many Australian Aboriginal crania (Lamach and Macintosh 1966, 1970). Therefore, the presence of a rudimentary styloid process, or its absence, may actually be evidence of evolutionary continuity within eastern Asia, although a left temporal from Xujiayao may have had a styloid process. If Asian Homo erectus was an ancestor of (at least some)

Homo sapiens crania then there must have been evolutionary reversals. Andrews (1984a), Stringer (1984a), and Wood

(1984) argued that there would have had to be too many evolutionary reversals and so this explanation was less parsimonious than one excluding the Asian Homo erectus material from the ancestry of anatomically modem Homo sapiens. This view is supported by the possibility that some of the Chinese Homo erectus material may chronologically overlap with archaic Homo sapiens crania from Europe and Africa, although there are problems with the dating of this material (cf. Chapter 7). If Hublin ( 1986) is correct, only one reversal is necessary - reduction in bony hypertrophy, especially of the posterior of the cranium. There is also a concentration of the features that differentiate the Asian Homo erectus material on the temporal bone and so these could possibly be functionally related, which would again require only one reversal.

account of individual variation, Stringer (1984a) adopted an approach where a range of characters is provided, the majority of which could be expected to be present on a member of a taxon. It is also uncertain if cladistics is applicable for assessing relationships between closely related species or subspecies or groups within a species, as is required for an assessment of the origins of anatomically modem humans (Habgood 1989c; Trinkaus 1990, 1992; but see Lieberman 1995). In an investigation of the origin of anatomically modem humans one is dealing with microevolutionary trends and not macroevolutionary speciation events and so when dealing with such anatomically and genetically similar populations, it is doubtful that any morphological differences between archaic and modem individuals/populations can be used to rule out relatedness. Kennedy ( 1991) observed that phylogenetic systematics lacks the resolution to be effective at infra-generic levels as it is at supra-generic levels. Also, Stringer observed ...Upper Pleistocene human evolution has so far received less attention from workers employing cladistic techniques, perhaps partly because of the difficulty of treating possible subspecific variation cladistically. (1987:136)

If Asian Homo erectus did not contribute to the origin of anatomically modem Homo sapiens, the presence within the Ngandong sample of sphenoids that do not contribute to the medial wall of the glenoid fossa, mastoid and supramastoid crests that unite anteriorly, higher cranial vaults and increased cranial capacity (Hublin 1986; Stringer 1984a) must also have been examples of parallelism. There is evidence of parallelism within hominid evolution such as the cranial capacity increase of the robust australopithecines. Lamach and Macintosh (1974) also identify three traits that they found to be unique to the Ngandong material. These are the location of the foramen ovale in a pit, the lack of a postglenoid tubercle and the positioning of the squamotympanic fissure along the floor of the glenoid fossa. These features cladistically separate the Ngandong hominids from not only African Homo erectus, but also earlier Asian material, yet, as discussed before, many other features link the Ngandong calvaria with Asian Homo erectus material (Santa Luca 1980; Stringer 1984a). Larnach and Macintosh (1974) did not find these features on any Australian crania examined and so concluded that if there was a direct evolutionary development from the N gandong material to Australian Aborigines these traits must have been "lost and their sites then reverted to a similar state to that of Homo erectus" (Lamach and Macintosh 1974:101).

Lieberman ( 1995: 160), whilst acknowledging that cladistic analyses have limitations when applied to closely related taxa, argued that this method was "a more logical technique for analysing the data used to support different theories about the origins of modern humans". However, because of the issues inherent in cladistics (Habgood 1989c; Trinkaus 1990, 1992), a phenetic approach has been adopted here (Holmes 1980; Pichardo 1978). A phenetic approach is interested in overall morphological similarity based on many characters, attempts to sample the total morphological variability evident within a sample and allows the grouping of individuals on the basis of overall morphological similarities. Thus, while the cladistic method emphasises discontinuities, the phenetic approach incorporates both similarities and differences. A phenetic approach approximates the "overall morphological pattern" of Le Gros Clark (1964). Trinkaus (1990) and Lieberman (1995) have suggested that a phenetic approach can ignore issues such as sample definition and trait identification. However, the phenetic approach used here has generally dealt with individuals and has utilised the concepts of primitive, shared, and derived morphological traits to assist in ascertaining evolutionary relationships between hominids because it is the combination of these types of traits that make up the unique total morphological pattern of populations.

The suggestion that Asian Homo erectus material is phylogenetically different to African Homo erectus and that only this latter taxonomic group contributed to the origin of anatomically modern Homo sapiens is difficult to justify on morphological grounds. Postcranial material also indicates that both African and Asian material displays an anatomical pattern that may be referred to as characteristic of Homo erectus (Day 1971, 1982, 1984; Kennedy 1983, 1983, 1984a, 1984b). A similar pattern is also evident on postcranial material from Arago and Castel di Guido, which are attributed to archaic Homo sapiens, suggesting an evolutionary link (Day 1982, 1984; Mallegni et al. 1983).

A debate developed between cladists and non-cladists. Halstead (1978:760) suggested "The cladists adhere to the tenets of Hennigism with religious fervor", while Charig argued that ...the virulence of the controversy .... comes almost entirely from the 'cladists' some of whom ...embrace their creed with a quasireligious fervor. Surely the subject of biological classification... merits a less emotional, more rational approach. (1981 :20)

From these examples it is evident that a major limitation of a cladistic approach is that it takes too narrow (presence/absence) a view and does not allow for morphological variation or effectively deal with shifting patterns of morphological variation. A consideration of morphological variation is necessary so as to fully understand the relationships of fossil and modern crania. To address the issue that a cladistic approach takes limited 11

not an issue for this study if Neanderthals and anatomically modem humans are different species (H. neanderthalensis & H. sapiens respectively), but rather what is at issue is their morphological similarities and differences. There will therefore, be no attempt to untangle the plethora of taxa that have been used for classifying Pleistocene hominids (i.e. H. ergastor, H. erectus, H. antecessor, H. georgicus, H. rhodesiensis, H. heidelbergensis, H. neanderthalensis, H. sapiens). The taxon Homo erectus will be used to encompass material that is grouped within the taxa Homo ergastor, Homo erectus sensu stricto and Homo erectus sensu lato and should be read as Homo erectus sensu lato. Where terms such as "Pithecanthropus" or "Sinanthropus" are used in reference to Indonesian or Chinese Homo erectus material it is not meant to infer that these redundant taxonomic na~es are valid.

It is not the intention of this study to add to this debate, but rather to explore the relationships between morphologically variable and closely related hominid groups, which may have a unique overall morphological pattern made up of morphological traits shared with other groups.

Methods and Data As noted earlier, the debate surrounding the ongm of anatomically modem Homo sapiens throughout the Old World is addressed in this study. Multivariate analyses and morphological comparisons are used to investigate when anatomically modem humans first appeared in sub-Saharan Africa, north Africa, western Asia, Europe, Australasia and east Asia in order to evaluate the two major explanations for the world-wide appearance of anatomically modem Homo sapiens . Willermet and Clark {1995) identified that researchers often use different data sets (measurements, morphological features, crania) to support their preferred modem human origins model or refute the opposing model and so are not using the same data to test the two hypotheses. In this study data presented to support each of the two hypotheses is assessed and evaluated, as opposed to a selective use of data. The following paragraphs outline the crania that have been studied and the data used and how it was obtained.

Only archaic Homo sapiens , Neanderthals and anatomically modem Homo sapiens crania have been analysed in detail. Homo erectus material from China and Indonesia were included in an earlier study (Habgood 1982, 1984b, 1988; Habgood and Walker 1986), in which they formed their own distinct group. The late Homo erectus sample from Ngandong was included in both the earlier study and the present investigation. In these studies the crania generally formed their own group (Chapter 6; Habgood 1982, 1984b, 1988; Habgood and Walker 1986). Also, other hominid material was considered when it was crucial to an understanding of the origin of modem humans in a particular region.

Crania and Taxonomy Due to the adoption of the phenetic approach and the concept of phyletic gradualism for the latter stages of human evolution, the recommendations of Mayr (1969) and Simpson ( 1961) have been followed. Simpson ( 1961) suggested that for the purposes of classification, lineages must be divided up, often arbitrarily, into distinct segments, while Mayr ( 1969) argued that due to breaks between horizons, species can be separated in a non-arbitrary way. Mayer's (1963) biological species concept has also been adopted. However, these recommendations are not universally accepted (for example, Eldredge and Gould 1972; Eldredge and Tattersall 1982; Gould and Eldredge 1977; Tattersall 1986; Wiley 1981).

Morphological definitions of Homo erectus, archaic Homo sapiens and anatomically modem Homo sapiens have been attempted (e.g. Andrews 1984a, 1986; Day and Stringer 1982; Jacob 1976; Howell 1978; Howells 1980, 1988; Lamach 1978; Lamach and Macintosh 1974; Le Gros Clark 1964; Macintosh and Lamach 1972; Santa Luca 1980; Stringer 1984a, 1985; Stringer et al. 1979; Stringer et al. 1984; Wood 1984). They have not however, been wholly successful (Aiello 1993; Bonde 1989; Brown 1990; Day and Stringer 1982; Foley 1989; Kidder et al. 1992; Lieberman et al. 2002; Tattersall 1986; Thome and Wolpoff 1991; Wolpoff 1986a, 1992a).

A combination of the concepts of chronospecies and palaeospecies has, therefore, been used to help define the broad group of crania to be studied (archaic and early anatomically modem Homo sapiens ), although this material does display a unifying suite of morphological features that binds it together and separates it from earlier hominids ( see also Smi~h et al. 1989; Stringer 1992). This is different to the evolutionary (phylogenetic) species concept of Stringer (1992), which relates to a lineage of ancestor-descendant populations and is popular among cladists (see Wiley 1981).

The defmition of "a primitive form of Homo sapiens" proposed by Stringer et al. (1979:248), along with the more general definition provided by Howells ( 1988), have been adopted here for archaic Homo sapiens. Where the term archaic modem is used it follows Howells (1988) and identifies anatomically modem Homo sapiens that are distinct from existing humans because they reserve a greater degree of robustness and ancestral/primitive traits such as unusually large brow-ridges and markedly prognathic faces. Neither of the terms archaic Homo sapiens or archaic modem are used to refer to Neanderthals. The Neanderthal definition provided by Stringer et al. (1984:Table 2) has been adopted and added to (Table 5:6). A number of adequate definitions of anatomically modem Homo sapiens are available which encompass the material that is here referred to this taxonomic group (Day and Stringer 1982; Lamach 1978:List 8; Lieberman et al. 2002; Le Gros Clark 1964; Stringer 1984a:Table 2, 1985:Fig. 2; Stringer et al. 1984:Table 1). Instead of using the phrase "anatomically modem humans" or "archaic Homo sapiens", one could use the terms "modem

It is admitted that there are problems with the application of the biological species concept to palaeontology and that chronospecies and palaeospecies (morphospecies) are arti~cial constructs with an unknown amount of biological reality. The evolutionary (phylogenetic) species concept also has problems in that, in its pristine form, it assumes that significant morphological change only occurs with speciation, whereas there is ample evidence of gradual phyletic evolution. However, as the aim of this study is not taxonomic classification, but rather an examination of morphometric relationships , these species concept issues do not create a methodological conundrum . For example , it is

12

populations" or "recent and modem populations" suggested by Wolpoff ( 1986a).

as

Steinheim, Feldhofer 1 and Arago 21, the originals were used to provide non-metric data, but some measurements were obtained from casts . Casts were also used for some measurements if an original cranium was deformed. This occurred with Steinheim, and Arago 21 where measurements from casts were used to supplement those from the originals.

The most contentious definition used is that of "archaic Homo sapiens". The validity of this group has been questioned by Tattersall (1986; see also Foley 1989), who argued that the Middle Pleistocene hominid sample that usually falls under this "umbrella label" was variable enough to be divided into at least three, or even more, species. Wolpoff and Caspari ( 1990) have also contested the validity of applying the taxonomic category of "anatomically modem Homo sapiens to Middle Palaeolithic (Europe & the Levant) or Middle Stone Age (sub-Saharan Africa) hominids.

Measurements Measurements have always been an important part of cranial analysis (Armelagos et al. 1982; Gould 1981). Univariate and multivariate analysis of cranial measurements became common practice during the first half of this century and continue to be practiced. The data used here consisted of neurocranial and splanchnocranial measurements of Middle and Upper Pleistocene hominids. The measurements were chosen because major structural changes occur in the face and the vault, and the face to calvaria relationship undergoes important changes during the development of anatomically modern humans. Using cranial data has other advantages, in that the skull (or parts thereof) is one of the most commonly preserved parts of the skeleton (Boaz and Behrensmeyer 1976), and displays more variation and change throughout the period under consideration than most other skeletal parts. Mandibular measurements were not used, as the mandibles of archaic Homo sapiens are less commonly preserved than the crania, and because the mandible is a very plastic bone and more open to external, non-evolutionary influences. Dental measurements were excluded because severe attrition (evident on many Middle and Upper Pleistocene crania) can significantly alter maximum dental measurements, especially mesio-distal length (Brothwell 1981; Frayer 1978; Wolpoff 197lc ), and because it has been demonstrated that "traditional" cranial measurements provide a better sorting criterion for human populations (Falk and Corruccini 1982).

The problem of whether the number of Middle Pleistocene hominid species has been under-estimated or not, does not affect this study because this sample is both chronologically and morphologically intermediate (in particular geographical regions at least) between Homo erectus and Homo sapiens. It is probable that some, at least, of this group must broadly represent the type that gave rise to anatomically modem humans. This is the question that is to be investigated here. It should also be remembered that by using the term archaic Homo sapiens to define a group of hominids does not necessarily imply that the group is a single species as the link between speciation and morphological variation is difficult, if not impossible, to demonstrate for this material (see discussion in Smith et al. 1990). Some of the crania used have been classified as Homo erectus by certain authors, but the overall morphological pattern would place this material within archaic Homo sapiens (Habgood 1989e). Data The use of measurements taken by different people using different measurement techniques and definitions can impact on the results of multivariate studies (Habgood 1984a, 1985a; Habgood and Walker 1986; Trinkaus 1984b). There are numerous studies of inter-observer error in cranial measurements and the impact this may have on (Jamison and Zegura 1974; Kemper and Pieters 1974; Page 1976; Utermoble and Zegura 1982). Most indicate that there may be significant differences when the same measurement of the same cranium is taken by different anthropometrists. For this reason, most of the measurements used were taken by the author on the original specimen or good casts of the hominid crania. A cast was regarded as "good" if the measurements obtained from it were not markedly different from the published measurements, which had been taken on the original specimen. Published measurements were only used when the original specimen (or a good cast of it) was not available. This occurred with hominids such as Shanidar 1, Dali and Eliye Springs 11693 (see Tables 2:1, 3:1, 4:1, 5:2, 6:1, 7:1).

At the beginning of this century, attempts were made to standardise cranial measurements so that different data could be compared. This attempt was not successful, as there remains three major sets of definitions of cranial measurements in use: those defined by Martin (Martin and Saller 1957), the Biometric School (for example Morant 1927, 1930), and Howells (1973c). Most of the measurements used in this investigation are those defined by Howells ( 1973c), with the addition of some defined by Martin (Martin and Saller 1957) and Smith (1976b). Table 1:1 presents the measurements that were used, the sources where detailed definitions of the measurements may be found, and the abbreviations of the measurements that have been used on the tables and scatter diagrams. It was found that with some of Howells' ( 1973c) measurements, such as minimum cranial breadth, which do not have well defined end-points, it was difficult to obtain equivalent values when the measurement was repeated, therefore these measurements were not used. Howells ( 1969a, 1973c) did not use arc measurements because he felt they were inefficient measures of shape. Arc measurements do, however, reflect the curvature of the vault, which may offer significant differences between archaic and modern crania (for example the receding frontal of Neanderthals and the high curved frontal of anatomically modem humans), hence various arc measurements were used in this investigation. Howells ( 1969a) also felt that the base of the skull was generally neglected when measurements have often been taken, and so a number of basicranial measurements

The hominid material studied dates from between approximately 600,000 years BP and 25,000 years BP, although some earlier and later remains were also considered when they were essential to ascertaining if anatomically modem humans developed in a region ( for example in east Asia and Australasia, where few hominids dating to this period have been found). The hominids examined for each region are listed in the respective chapters (Tables 2:1, 3:1, 4:1, 5:2, 6:1, 7:1). Only the original crania were used for determining non-metric data, whereas casts were adequate for mandibular non-metric data. In some instances, as with 13

Table 1:1. Cranial measurements used in analyses, their abbreviations and sources where they are defined. Measurement

Present Study

Howells 1973c

Martin and Saller 1957

Glabella-occipital length Glabella-inion length Nasion-occipital length Basion-nasion length Basion-bregma height Auricula-bregma height Maximum cranial breadth Maximum frontal breadth Minimum frontal breadth Bistephanic breadth Bizygomatic breadth Biauricular breadth Biasterionic breadth Basion-prosthion length Nasion-prosthion height Nasal height Nasal breadth Orbit height (left) Orbit breadth (left) Bijugal breadth Palate length Palate breadth (external) Palate corpus thickness at the border between the left first and second molars (Freedman and Wood 1977 CORP.B, No. 12) Premolar chord between the midpoints of first left premolar and second molar (Freedman and Wood 1977 Pl-M2 CH., No. 17) Prosthion-end of dental arch Temporal height (left) (Smith 1976b) Temporal breadth (left) (Smith 1976b) Bimaxillary breadth Bifrontal breadth Biorbital breadth (ectoconchion) Interorbital breadth (dacryon) Simotic chord Cheek height (left) Nasion-bregma chord (Frontal chord) Nasion-bregma arc (Frontal arc) Bregma-lambda chord (Parietal chord) Bregma -lambda arc (Parietal arc) Lambda _-opisthion chord (Occipital chord) Lambda-opisthion arc (Occipital arc) Lambda-inion chord Lambda-inion arc Inion-opisthion chord Nasion-opisthion arc Bimastoid breadth Mastoid-prosthion length (from left mastoid tip) Mastoid-nasion length (from left mastoid tip) Mastoid-lambda length (from left mastoid tip) Transverse arc via bregma

GOL GIL NOL BNL BBH ABH XCB XFB

GOL

I 2 Id 5

--

XCB XFB

-21 8 10

--LB H' OB B B" B'

WFB

--

STB ZYB AUB ASB BPL NPH NLH NLB OBH OBB JUB PAL PAB

STB ZYB AUB ASB BPL

--

45(1) 60

MAB

61

----

PCT

--

--

--

PLC

--

--

Morant 1936 m2pl

PDA TPH TPB ZMB FMB EKB DKB WNB

--

--

--

ZMB FMB EKB DKB WNB

----

--

49a 57

DC SC

WMH

WMH

--

--

FRC FRA PAC PAA

FRC

29 26 30 27 31 28 31(1) 28(1) 31(2) 25 13

SI ' SI S2' S2 S3' S3 /-I ARC/-1 I-O

occ

OCA LIC LIA IOC NOA MDL MPL MNL MLL TAA

14

-NOL BNL BBH

Biometric School Morant 1927 1930 L

NPH NLH NLB OBH OBB JUB

-PAC

--

occ ----

-----

----

9 !Ob 45 llb 12 40 48

-54 52

--

--

--24

-J

-Biasterionic B

-G'H NH NB O2L O'IL

--

--

s

--

---

--

Q'

were also recorded (Table 1: 1). The entire cranium was therefore, encompassed by the full suite of measurements. In some instances, especially for the splanchnocranium, where the cranium was damaged, bilateral symmetry was assumed so that a measurement could be estimated.

themselves). Also, if the same measurement correlates differently with other measurements in different cranial groups, the incorporation of two measurements in an index only complicates the multivariate analyses (especially Qmode methods) and interpretation of the results (Walker 1985). These problems are exaggerated when the same measurement is used in several indices. Atchley et al. (1976) have also used computer generated data to demonstrate the poor performance of pairs of variables x and y as the index x/y in multivariate analyses.

The maximum number of measurements used in any analysis was 46 (Tables 1:1, A:11, Figs. 5:9, 5:11). There were 16 skulls for which this full set of measurements was available (that is, a data matrix of 16 objects by 46 variables). This data set, which consisted of predominantly European archaic and anatomically modem crania ( including Kabwe 1 from sub-Saharan Africa and Skhiil 5 from western Asia), formed the basis for all the metrical analyses (Table 1:2). As the Gibraltar 1 cranium is damaged some measurements were estimated based on bilateral symmetry and compared to the measurements taken on a reconstructed cast. The Steinheim cranium suffered some postmortem warping and damage ( see discussion in Chapter 5). Measurements were therefore taken on both corrected and uncorrected casts of Steinheim held at the Natural History Museum, London.

The measurement procedures recommended Bowles (1974) were followed to ensure a relatively "accurate" and "precise" value for each measurement could be obtained. Bowles (1974) defined an "accurate measurement" as one that is in close agreement with the true "value", while a "precise measurement" is one which gives the same value when the same object is measured repeatedly . When the same original specimen was measured, there was found to be a high correspondence between measurements obtained in this study and those recorded by Stringer (pers. comm.).

Table 1:2. Crania that formed the basic data set for all metrical analyses and abbreviations for Figures and Tables (cf. Tables 2:1, 4:1 and 5:2 for sources of measurements). Crania Steinheim Kabwe 1 Gibraltar 1 La Chapelle-aux-Saints 1 La Ferrassie 1 Skhiil 5 Predmost 3 Predmost 4 Combe Capelle Cro-Magnon 1 Mladec 1 Chance lade Abri Pataud 1 Dolni Vestonice 3 Oberkassel 1 Oberkassel 2

Discontinuous-Discrete-Epigenetic-Non-metric Traits It has been argued that non-metric cranial traits may reflect genetic differences between populations more closely than measurements because they convey more genetic information about the individual and appear not to be influenced by skeletal plasticity (Berry 1976; Berry 1979; Berry and Berry 1967; Ossenberg 1976, 1977, 1981; Perizonius 1979; but see Corruccini 1976; Finnegan 1974, 1978; Lamach 1974b; Rightmire 1972).

Abbreviation St Kl G1 LCl LFl Sk5 P3 P4

However, there is evidence that the occurrence of some nonmetric traits are correlated highly with the occurrence of other non-metric traits (Hertzog 1968; Cheverud and Buikstra 1981b ), the frequency of some non-metric traits is influenced by cranial deformation (Anton and Weinstein 1999; Brown 1989; Ossenberg 1970), environmental stress may be important in controlling the frequency of bilateral non-metric traits (Pardoe 1986; Kennedy 1986; Trinkaus 1978d), and that there is considerable variation in the heritability of some non-metric traits (Cheverud and Buikstra 1981a).

cc

CMl Ml Ch APl DV3 01

02

Carpenter (1976) and Corruccini (1974b) found that when non-metric traits were used in isolation they had only limited discriminatory value, and felt they should be used only in conjunction with other osteological measurements and observations. Cheverud et al. (1979) and Corruccini (1976) have also demonstrated that some metric and some nonmetric traits may be highly correlated. Gruneberg ( 1963) went so far as to state that non-metric traits are basically metric variations. Sj0vold (1984) has also demonstrated that both cranial measurements and non-metric traits have high heritability.

Mean values were not used to replace missing data, and so the less complete crania were added to the basic data set and the variables, which the additional hominids could not provide, were eliminated from the data set. These modified data sets· were analysed separately. There would have been additional objects , but fewer variables in such data sets. As the number of variables became small (i.e. when fragmentary specimens were used) some groupings became less robust and some individuals more volatile. Only linear measurements were used because the combination of different mensuration scales (angles, areas or weights) can have deleterious effects on the precision of multivariate analyses (Anderson 1958; Kowalski 1972) and can make it difficult to interpret the components. Indices were not used because of their inherent disadvantages (Atchley et al. 1976; Corruccini 1978; Oxnard 1878, 1983 Rhoads and Trinkaus 1976). They can compound measurement error and increase the danger of swamping informative measures with less meaningful ones (the indices

It appears, therefore, that metric and non-metric characters may not be totally independent, either genetically or morphologically, and so the greatest amount of information in skeletal studies will be obtained when both forms of information are used in conjunction. For this reason, a limited number of cranial (Fig. 1: 1; Table 1:3) and mandibular (Table 1:4, Fig. 1:2) non-metric traits were analysed. Whilst the cranial non-metric trait assessment was 15

3

7

6

2 10

Figure 1 :1 Cranial non-metric traits used in this study.

16

Table 1 :3. Discontinuous, discrete, epigenetic, or non-metric traits used in study. CRANIAL TRAITS. For definitions see Berry and Berry (1967) for Nos. 1-7, de Villiers (1968) for No. 8 and Santa Luca (1978) for Nos. 9-11. 1. 2. 3. 4. 5. 6.

Ossicle at lambda present. Lambdoidal ossicles present. Parietal foramen present. Metopic suture present. Ossicle at asterion present. Supraorbital foramen complete.

CRANIA Steinheim

Kabwe Gibraltar 1 La Ferrassie La Chapelle 1 La QuinaH5 Spy I Spy2 Skhill 5 Jebel Qafzeh 9 Mladec 1 Cro-Magnon 1 Cro-Magnon 2 Abri Pataud I Oberkassel 1 Oberkassel 2 Engis I Laugerie Basse 4 Singa

1 0 0 1 0 1 1 0 0 1? 0 0 0 0 I 0 0 0 0 0

2 lL 0 IR 0 0 1B 0 0? lL 0 0 0 0 0? 0

IR 0 IL 0

7. Frontal notch or foramen present. 8. Inferior frontal eminence present. 9. Occipitomastoid crest present. 10. Occipital torus present. 11. Suprainiac fossa present.

NON-METRIC CRANIAL DATA TRAITS 3 4 5 6 7 0 0 lR? lR? 0 IL 0 0 0 1B 0? 0 0 lL lL 0* 0* 0 0? 0 1B 0 0 0 1B 0? IR 0 0 1B 0 0 0 0 1B 0 0 0 0 0 0 0 0 0 0 IR 0 0 IL 0 lL 0 0 0 1B lL IL IL 0 0 I?+ 0 0 I 1B 1B lR 0 0 0 lL 0 0 0 0 1B 0 0 I 0 0 0 0 0 1B 0 1B 0 0 0 IL 0 0 0 0

8 0 0 0 1 0 0 0 0 lR 0 1B

0 0 0 1B 1B

0 0? IL$

9 1# 1 IR 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0

10

1 1 1 1 1 1 I 1 1 0 0 0 0 0

Ix 0 0 0 0

11 1$ l? 1 1 I I I I 1$

0 0 0 0 0 lz 0 0 0 0

Key: L=left side; R=right side; B=both sides; ?=observation questionable due to poor preservation of relevant region; #=small; $=slight or shallow; x=large; z=deep; *= Santa Luca (1978) has drawn La Ferrassie I as though it has a lambdoid ossicle on the left side and an ossicle at the right asterion, but examination of the original did not reveal these, although there are breaks in these regions; +=this could be a break, but has been scored as a metopic suture.

presence/absence, the mandibular non-metric trait assessment was an ordinal partitioning of a continuous degree of expression (position, size of or degree of development) and so some specimens invariable fall near the arbitrary boundaries between the categories adopted, but as all observations and recordings were undertaken by the author a high degree of assessment consistency was been achieved. Forty non-metric cranial traits (defined by Berry and Berry 1967; Santa Luca 1978; de Villiers 1968) were originally recorded, although most were absent from the bulk of the crania examined. The eleven traits that were analysed were the only traits that could be found on at least one of the crania studied for which all eleven could be recorded. They do not form an exhaustive list of possible non-metric traits. Some of the traits used, especially those of the mandible, were chosen so as to differentiate between archaic and anatomically modem hominids.

results, can simultaneously analyse numerous objects and variables, provide simplified representation of the relationships between the objects and variables, and treat the objects (crania) as an integrated whole and not separate figures (Corruccini 1978; Doran and Hodson 1975; Howells 1969b; Sneath and Sokal 1973; Wright 1985). Numerous analyses of Middle and Upper Pleistocene hominids utilising various multivariate techniques have been conducted (e.g. Bilsborough 1972, 1973, 1976, 1984; Bilsborough and Wood 1986; Brauer 1980a, 1980b, 1981, 1984a, 1984b, 1984c; Brauer and Leakey 1986a, 1986b; Brothwell 1960; Corruccini 1974a, 1975b; Habgood 1988, 1984a, 1984b, 1985a, 1985b, 1986a; Habgood and Walker 1986; Henke 1984; Hills and Brothwell 1974; Howells 1970, 1975, 1983; Pietrusewsky 1979; Rightmire 1979; Stringer 1974a, 1974b, 1978; van Vark 1983a, 1983b, 1984). These references demonstrate that multivariate methods have a well-established part to play in palaeoanthropology. Sokal and Sneath (1963) have also suggested that multivariate morphometrics are especially useful at the taxonomic level of the species and below, which is how they have been used here.

Multivariate Techniques Multivariate techniques are of use because they clearly define what is being compared (measurement or contingency data), allow the potential for repeatability of the methods and 17

Tabl e 1 :4. Discontinuous, discrete, epigenetic, or non-metric mandibular traits used in this study.

1. Mental foramen number : 1 - single, 2 - multiple one side at least. 2. Mental foramen position: 1 - under PM3/PM4 , 2 - under PM4, 3 - under PM4/Ml , 4 - under Ml. 3. Retromolar gap: 1 - not present , 2 - present but small, 3 - present and large. 4. Development of a chin: 1 - no development-receding symphyseal region, 2 - slight development, 3 - well-developed. NON-METRIC MANDIBULAR DATA TRAITS MANDIBLES

No. 5

1 1- 60%

2 3- 60%

3 1- 60%

4 1- 80% 2-20%

European Early Neanderthal

7

2-71.5%

4-57%

1-43% 3-43%

1-86% 2- 14%

European "classic " Neanderthal

8

1- 50% 2- 50%

3- 50% 4-50%

2-37.5% 3- 50%

1-62.5% 2-37.5%

3-5

1-40% 2-40%

4-60%

2-40% 3-40%

2-60%

Border Cave

2

1- 50% 2- 50%

2- 100%

1- 100%

3- 100%

Klasies River Mouth No. 41815

1

2

2

1

3

Western Asian Neanderthals

5

1-20% 2- 80%

3-20% 4- 80%

3- 100%

1- 80% 2-20%

Tabiin C2

1

1

4

2

2

Skhul/Qafzeh

3

1-100%

2- 66% 4-33%

1-33% 2- 66%

3- 100%

Eastern European Upper Palaeolithic

7

1-100%

2-71.5 3- 28.5

1-71.5% 2-28.5%

3- 100%

Western European Upper Palaeolithic

6

1- 83% 2-17%

1-33% 2- 66% 3-33%

1- 100%

3- 100%

European Middle Palaeolithic

Vindija

Retromo/ar gap

J

Figure 1 :2 Mandibular non-metric traits used in this study.

18

Corruccini (1978) has outlined four criteria that morphometric analyses should meet if they are to be of use: 1) The utilisation of multivariate methods must be adequately justified. 2) The reasons for using particular data must be explained and the variables should be clearly defined. 3) Analysis of the data must be completely done with due consideration to the distinction between size and shape information and the results of several different methods should be considered. Adequate presentation of the results must also be provided. 4) The multivariate results must be related back to the original anatomy. Whilst not all analyses of skeletal material using multivariate methods have met Corruccini's ( 1978) four criteria, they have adopted for this study ..

standardisation procedure allows male, female and immature crania to be compared in the one analysis. To demonstrate the effects of the standardisation procedure, a brief example of its use will be outlined. Habgood ( 1985a, 1988) carried out three principal components analyses on a data set that consisted of nine published cranial measurements for each of seven immature and thirteen mature Middle and Upper Pleistocene hominids. The first analysis was carried out on the raw unstandardised data set. This analysis was dominated by the variable that accounted for the largest value for each of the crania (cranial capacity) and so the groupings evident from it were based on similarities of this single variable. The second analysis used a data set standardised by variable only (correlation matrix). It was dominated by the overall size of all variables and so grouped large (adult) skulls together and small (immature) skulls together. The third analysis was conducted on a double standardised data set. The groupings obtained from this analysis were based on the morphological (shape) similarities of the various crania, with immature Neanderthals (Le Moustier, La Quina H18 and Gibraltar-Devil's Tower) grouping with adult Neanderthals (La Chapelle 1, La Ferrassie 1, Guattari-Monte Circeo 1 and Shanidar 1), and immature and adult anatomically modem crania grouping together. Skhfil 1 and Skhfil 5 also grouped together in this analysis. This example demonstrates that dominance of size information can be reduced in a data matrix by utilising a double standardisation procedure.

Van Vark argued that ...a mere application of the elementary statistical techniques, as developed by mathematical statisticians, is not, in general, practicable; supplementary procedures are necessary, and the elementary techniques have to be modified by its users in an adaptive fashion... (1976: 112) Such modifications have been incorporated in the statistical package, used for this investigation, which was devised by R.V.S. Wright and E. Roper, formerly of the Department of Anthropology, University of Sydney, Australia. The package consists of four interrelated elements (a standardisation procedure, principal components analysis, sum of squares cluster analysis, and k-means cluster analysis) and two supplementary ordination procedures ( correspondence analysis and an ordered similarity matrix).

Principal Components Analysis The second element of the package consisted of the double standardised data matrix being put through an R-mode (portrays interrelationships between variables analysed) and Q-mode (portrays interrelationships between objects) principal components analysis (Andrews and Williams 1973; Blackith and Reyment 1971; Doran and Hodson 1975; Goodman 1972; Gower 1966; Hodson 1969; Jolliffe 1986; Joreskog et al. 1976; Kim 1975; Kowalski 1972; Lebart et al. 1984; Orloci 1978; Orton 1980; Pimental 1979; Sneath and Sokal 1973; Tabachrick and Fidell 1983; Wright 1985). Principal components analysis is one of the most widely used multivariate methods for the analysis of skeletal material. Pimental (1979:56) has stated "There is abundant evidence that biological interpretation of a PCA is possible".

Standardisation Procedure The first element of the package offers the opportunity for the double standardisation of the original data set of measurements of individual crania, not group means. The data matrix was standardised to remove overall size and to leave the morphologically (and taxonomically) more useful shape information (Campbell 1963; Corruccini 1973; Kowalski 1972). The standardisation procedure (subtract the mean and divide by the standard deviation) involved an initial standardisation by variable, which gives each variable equal weighting within the analysis and, in so doing, stops the large variables, such as glabella-occipital length, from swamping the smaller variables, such as nasal breadth. The single standardised data set was then standardised by object so as to remove all latent information about absolute size differences between the objects, leaving only shape information latent in the data matrix of z-scores (Corruccini 1973, 1975a, 1978). This standardisation procedure eliminated general components (a component with all signs the same), which are commonly regarded as reflecting absolute size (Bilsborough 1972, 1973, 1984; Campbell 1963; Corruccini 1973; Jolliffe 1986; Kowalski 1972; Oxnard 1978, 1983; Sneath and Sokal 1973), and left only bipolar components (a component with mixed signs). Gelvin (1983) has demonstrated that size information may be distributed throughout numerous components. Howells (1984) has also discussed this problem at some length. This

The major function of a principal components analysis is to reduce the dimensionality of a large data set while retaining the variation present in the original data set. Although covariances and correlations are not ignored, a principal components analysis concentrates on variances (Jolliffe 1986). It shows the similarity or dissimilarity between objects in a sample for which no parametric distribution need be assumed and so is useful for dealing with isolated fossil specimens. It attempts to describe the variation within the data matrix by calculating new sets of orthogonal (independent) axes, each of which includes progressively less of the total within-group variance. The first new axis or principal component accounts for the maximum variance in the sample, the second component is orthogonal to the first and accounts for the maximum remaining variance within the sample, and so on, until all of the variance within the data matrix has been accounted for. Each of the principal 19

while Wilmink and Uytterschaut (1984) stated that the computation of principal components prior to a cluster analysis was an accepted practice. These observations were implemented in this investigation in that the component scores obtained from the principal components analyses were used as the data matrix to be further analysed by the two separate clustering techniques, which formed the third and fourth elements of the statistical package used .

components is located by weighted combinations of all the original variables . The eigenvalues represent the percentage of the total withingroup variance accounted for by the associated components (cf. Table A: 11). The eigenvalue for a component is obtained by summing the eigenvectors (see below) for that component. When the eigenvalues are added together, the cumulative percentage of the total within-group variance accounted for by the various components is reached. This allows successive decreasing variation to be examined, and also allows the relative importance or explanatory power of a component to be evaluated. The percentage value for an eigenvalue , or group of eigenvalues, should not be regarded as the percentage of infonnation, because variance corresponding to the remaining components may be random "noise" (Lebart et al. 1984).

Some of the components with very low eigenvalues may contain "noise ", such as measurement error , and so their inclusion in further analyses could have a deleterious effect on the sensitivity of these analyses (Goodman 1972; Hardy and van Gerven 1976; Kowalski 1972). There are various methods available for removal of such residual component s (Jolliffe 1986 ; Kim 1975; Pimenta l 1979; Rummel 1970). For this investigation the discontinuity method was used to eliminate residual components (Rummel 1970). This method assumes that after the last meaningful or important component has been extracted , there will be a sharper drop or discontinuity in the eigenvalue s. The discontinuity is clearly evident when the eigenvalues are plotted. This method generally retained the component s that accounted for at least 70% of the total within-group variation .

A principal components analysis provides a diagrammatic summary of the relationships between the objects when the component scores are plotted on a bivariate graph (see Fig. 5:9, Table A:11). Such a plot of the first two components prov ides the "best" possible two-dimensional representation of the multivariate distance s between the objects . Distances between the objects on a bivariate graph reflect the objects ' individual responses to the variables on the components used for the graph. That is, if two objects are positioned close together on a bivariate graph, they have similar patterns of response to the variables , but if they are separated they have different responses to the variables.

An examination of reviews of clustering techniques reveals that no single way of dividing up a data matrix will, necessarily , be completely satisfactory, and so more than one clustering method should be utilised so as to verify the robustness of the groups identified (Doran and Hodson 1975; Everitt 1980; Hodson 1970; Orton 1980; Wilmink and Uytterschaut 1984). For this reason two clustering procedures were used to analyse the reduced data set. Clustering methods assume that groups exist within the data to be analysed, and so the purpose of applying them was to segregate the objects into distinct and clearly defined groups. It is also assumed that objects that are placed in the same group are more similar to each other than they are to members of other such groups .

A principal components analysis not only provides information about the relationships between objects, but, via the eigenvectors, it provides information on the contribution of each variable to the components (see Table A:l 1). The eigenvectors allow the identification of those variables which account for the greatest amount of variance on each component , and so strongly influence the placement of the objects. This is possible because the components are uncorrelated, and thus can be interpreted individually. The eigenvectors can be "read" so as to ascertain those variables that contribute most to each of the components and to establish the interplay between variables. The contribution of the variables to each component can also be plotted on bivariate graphs of the eigenvectors. Those variables that are positioned at either of the extreme ends (positive or negative) of the range of a component have a high loading on that component , and hence greatly contribute to the within -group variance accounted for by that particular component. They do, therefore, greatly influence the component scores of the objects . If two variables have a similar eigenvector value for a component , or appear close together on a bivariate graph, they may be interpreted as measuring the same thing in relation to the within-group variance. If a variable is positioned near the origin on a bivariate graph, it has a small loading on those two components and so contribute s little to the within -group variance accounted for by those components . The object and variable points belong to different hyperspaces and so cannot be placed on the same bivariate plot.

The first method used was a modified (by R.V.S. Wright and E. Roper) version of Orloci's (1978) SSA cluster analysis. This sum of squares cluster analysis (similar to Ward's Error SS method) is a polythetic, hierarchical and agglomerative method. It probes for structure within the matrix of component scores and, in doing so, reduces the number of units of study further by combining similar objects into clusters in the production of a dendrogram that then forms a new basis for interpretation. This modified clustering method deliberately weights the component scores in accordance with the magnitude of the original eigenvalues (R.V.S. Wright pers. comm.) and uses a scale of distance to form the dendrogram based on the square root of the sum of squares of the members of each cluster (scaled between Oand 100 by the formula : range of the sum of squares minus the sum of squares for each cluster divided by I% of the range). The clusters are formed by the sequential addition of individuals. A "good" cluster is one where all of the objects unite at an early stage on the dendrogram and which does not join another cluster until well up the dendrogram . It should be stated clearly that a dendrogram is not a cladogram and no phylogeny is necessarily implied by the groupings evident on a dendrogram, although they may have some phylogenetic significance (Morton and Lalouel 1973).

Clustering Techniques Tabachrick and Fidell (1983 :397) observed that as the components from a PCA are orthogonal , "their use in other analyses may greatly facilita te interpretation of results", 20

In this study correspondence analysis was used to analyse both contingency tables (non-metric cranial and mandibular traits) and cranial measurements. In all analyses the raw data set was transformed so that the variables were given equal weight and the first two axes were plotted. It was found that there was a very strong similarity between the results obtained when a data set was analysed by either principal components analysis or correspondence analysis (compare Figs. 5:9, 5:11 & Table A:11).

The second clustering technique used was the k-means method (Doran and Hodson 1975; Everitt 1980; Gordon 1981; Hodson 1970; Orton 1980; Wilmink and Uytterschaut 1985). The same data matrices (reduced component scores from the principal components analysis) as used for the sum of squares clustering method were used for analysis by the kmeans technique. This is a non-hierarchical method that determines the "best" number of clusters (k) for the data matrix being analysed and assigns the objects to these clusters. The various clusterings are done independently of each other, thus two objects which are in the same group when the sample is partitioned into four clusters, may not necessarily be in the same group when the sample is partitioned into three clusters (Orton 1980). To find the "best" number of groups, the percentage error of fit (the averaged squared distance of the objects from the centres of their respective cluster, expressed as a percentage of their average squared distance from the centre of the whole group of objects thought of as a single cluster) for the respective clusters is plotted on a graph. The "best" number of clusters (if one exists) will precede a marked elbow or discontinuity in this graph (Orton 1980). The k-means program used here produced ten clusters, giving a greater partition of the data than was generally needed. On occasions the k-means method produces a "local", rather than a "global", optimum solution. It is, therefore, advisable to run the program a number of times, and, if there are different assignments of objects between the runs, the lowest percentage error of fit for the cluster, where it occurs between the various runs, should be used for the plot (R.V.S. Wright pers. comm.). Each data matrix was analysed by the k-means method on three separate occasions so as to obtain a "global" optimum. However, few "local" solutions were obtained in this investigation.

In general, the groupings identified by the four methods (principal components analysis, sum of squares cluster analysis, k-means cluster analysis and correspondence analysis) were very consistent and so the clusters are very "robust", which suggests that they are "real" and meaningful. Ordered Similarity Matrix The final ordination method used was developed by Pielou (1979, 1983). The ordered similarity matrix contains a similarity index for every possible pair of entities in the data set. It allows a calculation of a proportional similarity index, which eliminates factors of absolute size in the data, although the variables will not contribute to the analysis equally (R.V.S. Wright pers. comm.). The appearance of the matrix demonstrates whether there is gradual or discontinuous change through the sequence under study. This method was only used for western Asia (Table 4:7), Europe (Table 5:8) and Australasia (Table 6:3). Although the time intervals between the crania in the various analyses were not the same, as recommended by Pielou ( 1979), the crania were placed in a relative chronological sequence and so the analyses should still be valid (R.V.S. Wright pers. comm.). When the ratio Q/Q max, which records the disarray of the matrix (Q being the grading index and Qm the maximum value of Q, cf. Pielou 1979, 1983 for further details), was plotted, none of the three ordered similarities were shown to display gradual or continuous change throughout the sequence, with all being close to the border between ill-graded and well-graded matrices (Pielou 1983:Fig. 6).

Wilmink and Uytterschaut (1984), who evaluated various clustering techniques, found that component scores clustered by Ward's method and the k-means method produced "robust" clusters. Two other ordination techniques were also used to analyse the various data sets.

Morphometric Comparisons Wright (1985) observed that multivariate methods were not often used as heuristic devices that help guide further research. In this investigation this has been done, in so far as the results obtained from the multivariate methods were compared with the morphological characteristics of each individual hominid and were used to help elucidate relationships between hominids. The multivariate results and morphological descriptions based on personal observations made on original specimens or casts, examination of photographs and from published descriptions were used to evaluate the various explanations for the origin of anatomically modem humans in both regional and worldwide contexts.

Correspondence Analysis Correspondence analysis is an ordination technique similar to principal components analysis (B0lviken et al. 1982; Gauch 1982; Hill 1974; Jolliffe 1986; Joreskog et al 1976; Lebart et al. 1984; Lessertisseur et al. 1974; Teil 1975; Wright 1985). Although correspondence analysis is generally preferred when the data are contingency tables, it is also applicable to measurements (Hill 1974; Joreskog et al. 1976; Lessertisseur et al. 1974; Wright 1985). Correspondence analysis combines the advantages of both R-mode and Q-mode techniques in that it analyses the objects and variables together in the same low-dimensional hyperspace. This means that, unlike principal components analysis, correspondence analysis provides a representation of both the objects and variables on the same bivariate plot (see Fig. 5: 11). This allows the "correspondence" between objects and variables to be studied simultaneously. The conventions used to interpret a bivariate plot from a principal components analysis are also applicable to bivariate plots from correspondence analyses. Suffice it to say that an object is located near those variables that are similarly weighted (best associated) on the components plotted, and vice versa.

Most investigations of the complex problem of the origin of modem humans have dealt with the hominid sequences in particular geographical regions (cf. Smith and Spencer 1984 and references therein), although some have addressed the world-wide phenomenon of the appearance of anatomically modem humans by at least 40,000 years BP (Frayer et al. 1993; Smith 1985; Smith et al. 1989; Trinkaus 1982a; Wolpoff 1980a). The analyses of archaic Homo sapiens and early anatomically modem Homo sapiens crania from around 21

each of the geographical regions are dealt with in relative chronological order, and their dates, morphologies and positions within the multivariate analyses (if they were included) are discussed . The various explanations for the origin of modem humans in each region are evaluated in the light of the morphological and multivariate analyses. Ultimately the inferences drawn from the results of the analyses of the skeletal samples from the separate geographical regions are drawn together in order to ascertain when and where anatomically modem Homo sapiens evolved.

the world undertaken as part of this study attempts to shed light on the problem of the origin(s) of anatomically modem humans and to evaluate the applicability of the two prevailing hypotheses - the Multiregional Hypothesis and the Replacement Hypothesis . The Old World has been divided up into six major geographical regions: sub-Saharan Africa, north Africa , western Asia, Europe , Australasia and east Asia. Each geographical region is assessed separately and its skeletal sample evaluated independently in order to ascertain if anatomically modem humans developed locally or migrated into the area. The major archaic Homo sapiens crania from

22

CHAPTER TWO. SUB-SAHARAN AFRICA This revised chronological framework and the enlarged sample of hominid material will form the basis of this investigation (Figs. 2: 1, 2:2). A number of major surveys of the sub-Saharan African late Middle and early Upper Pleistocene hominid material have been published (Brauer 1984a, 1984b, 1984c, 1989a, 1989b, 1992; Rightmire 1984a, 1989). These surveys contain chronological and morphological discussions of the material that will be dealt with here, and some additional more fragmentary remains that will not be considered in any detail (Figs. 2:1, 2:2, Table 2:1).

Introduction This chapter will discuss hominid material recovered from sites throughout sub-Saharan Africa dated to the Middle and early Upper Pleistocene period. Late Homo erectus material from east Africa dating to approximately 1myr appears to document the appearance of more modem morphological features (Asfaw et al. 2002; Abbate et al. 1998). The Buia cranium from Eritrea has a suite of morphological features typical of Homo erectus including a long ovoid braincase, small endocranial capacity, greatest cranial breadth at the supramastoid crests and a large supraorbital torus, coupled with a more evolved feature in that the parietals reveal a high position for greatest biparietal breadth (Abbate et al. 1998). A similar mosaic of features is evident on the Daka calvaria from Ethiopia (Asfaw et al. 2002). These hominids provide a morphological link between earlier Homo erectus material and later Middle Pleistocene hominids in Africa.

The completeness of the sub-Saharan African late Early and Middle Stone Age hominid material varies greatly from isolated teeth and mandibular fragments such as those from Mumba Rock Shelter, Tanzania, and Equus Cave, South Africa (Brauer 1984a; Grine and Klein 1985), to almost complete crania such as that from Kabwe in Zambia. For this reason it was only possible to include sixteen hominids in the analyses carried out (Table 2: 1). Fragmentary material, such as that from the South African sites of Border Cave and Klasies River Mouth, which is crucial to an understanding of hominid evolution in the region, will also be dealt with. The hominids will be discussed in approximate chronological order. Further anatomical details for a number of the hominids are discussed in Chapters 6 and 7, and are presented in Tables 6:5 and 7:3.

The importance of sub-Saharan Africa in the study of the origin of anatomically modem humans has significantly changed during the last fifty years. This has been due to a major revision of the chronology of the African Stone Age, the discovery and study of new hominid material and genetic studies (dealt with in later chapters). During the 1950s radiocarbon dates suggested that the African Middle Stone Age was contemporary with the European Upper Palaeolithic and dated between approximately 40,000 and 10,000 years BP (Clark 1959). Since 1972 a revised chronology has indicated that the Middle Stone Age is contemporary with the European Middle Palaeolithic with many sites dating in excess of 100,000 years BP (Vogel and Beaumont 1972). Potassium-argon dates from Lake Ziway, central Ethiopia, indicated that the Middle Stone Age of east Africa, at least, began prior to 180,000 years BP (Wendorf et al. 1975), while lithostratigraphic analyses at Border Cave have been interpreted as indicating that the Middle Stone Age there began around 195,000 years BP (Butzer et al. 1978).

Bodo D'Ar An almost complete face and partial neurocranium, a left parietal fragment from a second individual and a distal portion of a humerus of a third individual were found scattered across a conglomerate and sand deposit near the Awash River in Ethiopia (Fig. 2:1; Asfaw 1983; Clark et al. 1994; Conroy et al. 1978). The large and robust Bodo 1 consists of most of the face and frontal, portions of both parietals and part of the right side of the occipital squama (Plate 1; Asfaw 1983; Conroy 1980; Rightmire 1996). The supraorbital torus is large and the glabella is prominent. The facial skeleton is massively constructed, especially the malar bone and the wide interorbital region. The zygomatics shelve evenly onto the maxillae. The nasal root is flattened and appears to have a relatively horizontal frontonasal suture. The large nasal aperture is very broad. There are no canine fossae and the maxillary sinuses are extensive. The palate is broad and deep. There is marked facial prognathism and postorbital constriction. The thick frontal bone is low and receding and displays slight sagittal keeling and a bregmatic eminence, while the parietal and occipital fragments indicate that the vault may have been short and rounded. Striations on the parietal suggest that the upper border of the temporal bone would have followed a high arched course (Rightmire 1996). Estimates of the cranial capacity of Bodo 1 range between 1300cc and 1500cc, with the lower figure more probable (Rightmire 1996). There are also what appear to be cut

This revision in the dating of the Middle Stone Age meant that Africa could no longer be seen as culturally lagging behind Europe and has also moved the Early and Middle Stone Age associated hominid material back into the late Middle Pleistocene. Many new late Early and Middle Stone Age hominid remains have also been recovered and published during the last few decades while others have been re-interpreted (Andrews and Stringer 1989; Beaumont 1980; Beaumont et al. 1978; Brauer 1984a, 1984b, 1984c, 1989a, 1989b; Brauer and Leakey 1986a, 1986b; Brauer and Singer 1996; Brauer et al. 1992; Churchill et al. 1996; Clarke 1976, 1985, 1990; Conroy 1980; Conroy et al. 1978; Day 1969, 1972; Day and Stringer 1982; Day et al. 1980; Deacon 1989; Grine and Klein 1985; Grine et al. 1998; Magori and Day 1983; Mehlman 1984, 1986; Morris 1992; Pearson, and Grine, 1997, Protsch 1975, 1976b, 1981; Rightmire 1975, 1976, 1978a, 1978b, 1979, 1981b, 1983; 1984a, 1984b, 1989, 1996; Rightmire and Deacon 1991; Singer and Wymer 1982; de Villiers 1973, 1976; White et al. 2003)

23

Table 2:1. Sub-Saharan African crania examined, abbreviations for Figures and Tables, and source of data. Hominid Border Cave 1 Border Cave 2 Border Cave 3 Border Cave 5 Bodo 1 Bodo BOD-VP-1/1 Boskop Byneskranskop material Cape Flats Cave of Hearths Eliye Springs KNM-ES 11693 Eyasi 1 Eyasi 2 Fish Hoek 1 (Skildergat or Peers' Cave) Florisbad Herto Bouri material Kabwe 1 Kabwe 2 Klasies River Mouth KRM 41815 (and other material) KNM-ER3733

Orieinal or Cast Original Original Published Original Cast Published Original Originals Original Original Published Cast Cast Original Original Published Original Original Originals, Published and Casts Cast

KNM-ER3883

Cast

KNM-ER 3884 (Ileret) Laetoli H18 (Ngaloba Beds) Matjes River material Ndutu Oakhurst material Olduvai H9 Omo (Kibish) 1 Omo (Kibish) 2 Saldanha 1 Saldanha 2 Singa Springbok Flats (Tuinplaas)

Published Cast Originals Cast Originals Original Cast Cast Original Original Original Original

Abbreviations

-BC2

-BC5

--

---

---ES

----

--Kl

-KRM41815

---

-Ll8

-N

---Om2

--Si

--

marks on the face and vault of Bodo 1 indicating possible defleshing of the cranium (White 1986).

Principal Source of Data Original Original De Villiers 1973 De Villiers 1976 Cast Asfaw 1983 Original Originals Original Original Brauer and Leakey 1986, 1986b Cast Cast Original Original White et al. 2003 Original Original Casts, Rightmire and Deacon 1991; Singer and Wymer 1982 Cast, Rightmire 1984, Leakey and Walker 1985 Cast, Rightmire 1984, Leakey and Walker 1985 Brauer et al. 1992 Cast Originals Cast Originals Original and Cast Cast Cast Original Original Original Original

deposits are comparable to those from the upper members of the Olorgesailie formation in Kenya (Members 10 and 14), which are dated to between 500,000 and 700,000 years BP (Clark et al. 1994). Radiometric ages (argon-argon) and biostratigraphy have been used to suggest an age of approximately 600,000 years BP for the Bodo hominids (Clark et al. 1994).

The second individual, BOD-VP-1/1, is represented by the robust posteroinferior comer of a left parietal (Asfaw 1983). It displays a large angular torus, a prominent supramastoid crest extension, a well-pronounced angular sulcus and very thick diploe. As with Bodo 1 the upper border of the temporal bone may have followed a high arched course (Rightmire 1996).

There is, however, the possibility that the Bodo cranium, at least, may be in a reworked association with the Acheulean artefacts and fauna! remains (White 1986).

The humerus fragment (BOD-VP-1/2), which consists of the distal segment lacking an articular portion, is described as relatively small with some cortical thickening (Clark et al. 1994). The differences in size and robustness between this fragment and the cranial material may be due to sexual dimorphism, with the former being female and the latter male (Clark et al. 1994).

The Bodo 1 cranium was found to group with Kabwe 1 and then the European fossils Petralona and Arago 21. This grouping reflects strong loadings on the variables for the breadth of the midfacial region and the form of the frontal squama. Brauer ( 1984a, 1984b) found a similar relationship between Bodo 1, Petralona and Kabwe 1 in a principal components analysis of facial data.

The three hominids are thought to have weathered from the lower Bodo Member of the Wehaietu Formation, which on palaeontological and archaeological grounds, was attributed to the first half of the Middle Pleistocene (Asfaw 1983; Comoy et al. 1978; Comoy 1980; Kalb et al. 1982a, 1982b, 1984). The archaeological and faunal assemblages from these

These groupings are consistent with the morphological affinities of Bodo 1. Petralona and Arago 21 are similar to Bodo in their degree of robustness and overall morphology such as supraorbital torus development , large dimensions of the nasal aperture and interorbital region, robust malars ,

24

Wadi

Haifa •

•

Jebel

Sahaba

Singa •

Eliye

• omo 1 .lleret

Springs.,

():Nctut _u Laetoli

\

.evas

Mu,;,ba

~

Kabwe. •Mumbwa

Cave of. Hearths

Figure 2:1. Map showing Sub-Saharan African sites mentioned in text.

25

Age in kyrs B.P.

90

dJ 1B

?

100

~♦

♦

? BORDER CAVE

~~ KLASIES RIVER

MOUTH1A

[)cb

LAETOLI~ H18 ~

0

SINGA

HERTO

200 . ~EYASI

1

~

~

FLORISBAD ILERET

300 KABWE 1

400

q

D

NDUTU

600

SALDANHA 1

0

BODO

Figure 2:2. The chronological position of the Sub-Saharan African hominid sample.

26

presence of an angular torus (Arago 47) and the degree of facial prognathism and postorbital constriction. Kabwe 1 is also quite similar to Bodo 1 except that it is not as robust. The shorter face of Bodo 1 is even more prognathic and its midfacial region is broader and flatter. In frontal morphology, supraorbital torus form, lack of canine fossae and degree of postorbital constriction they are similar.

Ndutu Beds and so the age bracket could be 200-400,000 years BP. Clarke ( 1990) also documents the possible association of Upper Acheulian handaxes with the hominid (from the same deposits) that date between 300-500,000 years BP. McBrearty and Brooks (2000) cite preferred age estimates of 490-780,000 years BP (palaeomagnetism) and 370-990,000 years BP (argon-argon ratio and palaeomagnetism) for Ndutu. The dating of the Ndutu hominid remains problematical.

On a principal components analysis of the splanchnocranium, Brauer (1984a) found Bodo 1 and Petralona to fall near each other, with both being separated on one axis from Kabwe 1. In overall robustness and degree of postorbital constriction, Bodo 1 is also similar to Saldanha 1, but they differ in that Bodo 1 has lower frontal and a supraorbital torus that does not taper laterally. Like Bodo 1, Eliye Springs 11693 and Florisbad also have quite broad faces.

Ndutu clustered with Kabwe 1 (the only sub-Saharan African hominid in the analyses), before joining with archaic Homo sapiens from Europe (Fig. 2:3, Table 2:2). On the correspondence analysis (Fig. 2:4) Ndutu fell on its own. The Table 2:2. K-means cluster analysis with Ndutu. Analysis includes 76.8% of the total within-group variance.

Rightmire ( 1996) details morphological similarities and differences between Bodo 1 and Homo erectus material and groups it with other Middle Pleistocene hominids from Africa and Europe including Kabwe, Elandsfontein, N dutu, Arago and Petralona. Stringer et al. (1979) placed Bodo 1, along with Kabwe, in their Homo sapiens Grade 1. Brauer (1984a) placed Bodo 1 in his Homo sapiens Grade 1 (early archaic Homo sapiens ), along with Kabwe 1, Saldanha, Ndutu, Eyasi, Cave of Hearths and some of the north African Homo erectus material. McBrearty and Brooks (2000) placed Bodo, along with Ndutu, Kabwe, Saldanha and Homo erectus material into their Group 1.

Group 1 Steinheim, Kabwe 1, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Ndutu. Group 2 Cro-Magnon 1, Chancelade, Dolni Vestonice 3. Group 3 Skhiil 5, Combe Capelle, Oberkassel 1, Oberkassel 2. Group 4 Predmost 3, Predmost 4, Mladec 1, Abri Pataud 1. variables that document the form of the occipital squama and the breadth of the cranial vault had high loadings (Fig. 2:4, Table A: 1). Andrews placed Ndutu within a multivariate analysis (cf Andrews and Williams 1973) and found it to group with Zhoukoudian Homo erectus material (quoted in Clarke 1976).

Ndutu A very fragmentary human cranium was found near the northern shore of Lake Ndutu at the west end of the main gorge at Olduvai, Tanzania (Fig. 2:1; Mturi 1976). The fragmentary cranium (Clarke 1976, 1990; Rightmire 1983) consists of the posterior parietals, both temporals, most of the occipital and portions of the frontal and facial segments. The cranium is relatively small despite the robust nature of some of the individual bones. There is marked postorbital constriction and indications of facial prognathism. There appears to have been some degree of parietal bossing and a lack of mid-sagittal keeling. The maximum cranial breadth is at the supramastoid crests. The occipital bone is thick and strongly flexed. The almost vertical upper scale is larger than the nuch~l segment. There is a well-developed occipital torus that is most prominent medially, and which has a flattened surface. Above this central portion there is a supratoral sulcus. From the preserved facial and frontal fragments it appears that the brow-ridges would have been welldeveloped, the interorbital region broad and the mid-face relatively gracile. There is also the possibility of the presence of a slight canine fossae (Brauer 1984a).

Morphologically Ndutu is similar to Kabwe 1, especially in the shape of the occipital squama and in details of the mastoid region (Rightmire 1983). Ndutu lacks the midsagittal keeling of Kabwe 1 and has a greater degree of parietal bossing, which is more reminiscent of Saldanha 1. The flattened surface of the occipital torus is matched by that on Omo 2 (Rightmire 1983). A reduced nuchal plane and expanded occiput can also be seen on Eliye Springs 11693, Omo 1 and Laetoli H18. A low position for maximum cranial breadth is also seen on Saldanha 1, Kabwe 1, Eliye Springs 11693 and Omo 1. The slight canine fossa is similar, although not as well marked, to that found on Kabwe 2, Eliye Springs 11693, Laetoli H18 and Florisbad. Clarke (1990) identifies similarities between Ndutu and Sale calvaria, including overall size and shape, degree of parietal bossing, lack of parietal keeling and supramastoid morphology. The differences between Ndutu, Bodo 1, Saldanha 1, and Kabwe 1 may be due to sexual dimorphism, with the smaller Ndutu cranium being female, while the other three are male.

The Ndutu find is currently dated to the later Middle Pleistocene, generally around 400,000 years BP, on the basis of a suggested correlation of the tuff that overlies the sandy clay from which it is derived with the Norkilili Member of the upper Masek Beds at Olduvai Gorge (Mturi 1976; Leakey and Hay 1982). However, Clarke (1990) contends that mineralogical analysis cannot conclusively differentiate between the tuffs of the Upper Masek Beds and overlying

Clarke ( 1990) details morphological similarities and differences between Ndutu and Homo erectus material and assigns it to archaic Homo sapiens. Stringer et al. (1979) placed Ndutu, along with Saldanha and Eyasi, in their Homo sapiens Grade 1 or 2. Brauer ( 1984a) placed Ndutu, along with Bodo 1, Kabwe 1, and Saldanha, in his Homo sapiens

27

0

100

50

St

Kl N

Gl LCl LFl SkS 01

cc 02 P3 P4

Ml APl CMl

Ch DV3

Figure 2:3. Dendrogram from sum of squares cluster analysis with the Ndutu cranium. A2

.Kl Ass·

• Ml

.01

·cc

.st .Sk5

OCA.

A Pt•

0

·02

N.

•p3 0

Ch•

.occ • p4

·c,

GOL

·ov3

LFl. •Lei

·xce

-1 0

XFB

.CMl -2

(-3.1)

I

A1 -1

0

Figure 2:4. Correspondence analysis with Ndutu.

28

Grade 1 (early archaic Homo sapiens). McBrearty and Brooks (2000) placed Ndutu, along with Bodo, Kabwe, Saldanha and Homo erectus material into their Group 1.

Saldanha-Elandsfontein-Hopefield The hominid remains from Saldanha were surface finds found on a deflation zone in the Hopefield District of the western Cape coast, South Africa (Fig. 2:1; Drennan 1953a, 1953b). Saldanha 1 (Drennan 1953a, 1953b, 1954; Singer 1954) is a large and robust calvaria displaying marked postorbital constriction, a low receding frontal with a massive supraorbital torus and an angled occipital squama with a medially prominent occipital torus. The parietals are gently rounded with only very slight sagittal keeling. The greatest cranial breadth would have been low on the vault towards the missing mastoid region.

those of Corruccini (1974a). On his principal components analysis of the frontal squama, Brauer (1984a) found a similar relationship between Saldanha 1 and Kabwe , but not with Omo 1 (Omo 2 was not included). Laetoli HIS which I did not include in an analysis with Saldanha 1) also fell near Saldanha 1 on Brauer's (1984a) analysis. On Brauer's (1984a) principal components analysis of the parietals, Saldanha did not cluster with any archaic hominids, but Kabwe 1 and Laetoli H 18 were moderately close to it. Table 2:3. K-means cluster analysis with Singa, Omo 1, Omo 2 and Saldanha 1. Analysis includes S3.2% of the total within-group variance. Group 1 Predmost 3, Predmost 4, Cro-Magnon 1, Dolni Vestonice 3, Chance lade. Group 2 Mladec 1, Abri Pataud 1. Group 3 Skhiil 5, Combe Capelle, Oberkassel 1, Oberkassel 2.

Saldanha 2 (Drennan and Singer 1955) consists of part of the right mandibular ramus. This fragment had a similar degree of fossilisation to the main vault bones of Saldanha 1 which were recovered 4.5 metres away, but differed in colour. Small pieces of parietal that joined Saldanha 1 were found near Saldanha 2 and Drennan and Singer (1955:364) concluded that "it [the ramus fragment] probably belonged to Saldanha Man". The size and shape of the fragment and reconstructions of it suggest that the ramus was broad and that the sigmoid notch was shallow (Drennan 1955; Drennan and Singer 1955). On the lingual surface there is the impression of the posterior root socket of the third molar, indicating that there was no retromolar gap.

Group 4 Steinheim, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Singa. Group 5 Kabwe 1, Omo 1, Omo 2, Saldanha 1.

Morphologically Saldanha 1 is very similar to Kabwe 1 in the degree of robustness and postorbital constriction and in the size and overall form of the cranium. They differ in that Saldanha 1 has a supraorbital torus that tapers laterally, less occipital protrusion, a less horizontal nuchal plane, a reduced occipital torus, no sagittal keeling and parietals that display more bossing and lack postbregmatic and pre-lambdoid flattening. Omo 2 has a similarly flexed occipital squama to Saldanha 1, but with a more laterally developed occipital torus. Omo 1 only displays superficial resemblances to Saldanha 1 in that it is less robust and more modem in appearance. It is more similar to Omo 2 and Laetoli HIS than to Saldanha 1.

As the Saldanha hominid remains were surface finds their dating is problematic. Their association with the faunal remains and Acheulean artefacts from the site is not secure because they were also predominantly surface collections. One excavation has documented the association between Acheulean artefacts and fauna (Singer and Wymer 1968), while uranium and fluorine analyses have suggested that the hominid remains and the faunal material from the site were essentially of one age (Oakley et al. 1977). Other studies, however, indicated that the faunal assemblage may have accumulated over a long period of time (Hendey 1974; Vrba 1982). The faunal assemblage is archaic in nature (Butzer et al. 1974; Howell 1978; Klein 1973b) and clearly dates to the Middle Pleistocene. Vrba ( 1982) suggested that there were two faunal components in the assemblage, one older than the hominid remains and a later mid-Pleistocene element that is probably associated with the hominid( s) · and dates to between 600,000 and 300,000 years BP. The morphology of the hominid material suggests a date towards the younger end of this range. McBrearty and Brooks (2000) cite a preferred age estimate of 780,000-1.2 m.y.a. for the Saldanha hominids based on associated fauna.

Three other fossils that were not included in analyses with Saldanha 1, but which are morphologically similar to it are Ndutu, Eyasi 1 and Eliye Springs 11693. The Ndutu cranium is much smaller than Saldanha 1, but is similar in its degree of postorbital constriction, lateral thinning of the occipital torus, supraorbital torus form, a lack of mid-sagittal keeling, a low position of maximum cranial breadth and parietal bossing, although expressed to a greater extent. The fragmentary Eyasi 1 calotte, which is also smaller than Saldanha 1, is similar in cranial contour, especially of the frontal squama, degree of postorbital constriction and a medially developed occipital torus, although it is not as developed. Eliye Springs 11693 displays some similarity to Saldanha 1 in frontal curvature and the low position of maximum cranial breadth.

Saldanha 1 was found to group with Kabwe 1 and then Omo 1 and 2 (Table 2:3). These relationships reflect strong loadings on the variables that document the mid-sagittal profile of the crania . In earlier analyses Saldanha 1 was also found to group with Kabwe 1 (Habgood 1982, 1984b; Habgood and Walker 1986). The results are consistent with

Stringer et al. ( 1979) placed Saldanha, along with Ndutu and Eyasi, within their Homo sapiens Grade 1 or 2, while Brauer placed it, along with Bodo 1, Kabwe 1, Ndutu and Eyasi, in his Homo sapiens Grade 1. McBrearty and Brooks (2000)

29

placed Saldanha, along with Bodo, Ndutu , Kabwe and Homo erectus material into their Group 1.

being a male and Kabwe 2 being a female, although the chronological association of the two is questionable (Stringer 1986).

Saldanha 2 is similar to the mandible from Mauer, Germany, in having a broad ramus , a shallow sigmoid notch and in the height and curvature of its anterior border (Drennan and Singer 1955). This morphological pattern is quite different from the earlier Homo erectus mandibles from north and east Africa, although there is a slight resemblance to the almost complete mandible from Lake Baringo, Kenya, which also has a broad ramus (Rightmire 1980). A proposed reconstruction of Saldanha 2 indicates that it would have been too short and broad to have fitted Kabwe 1 (Drennan 1955).

Most of the postcranial material , which is from at least three individuals (Oakley et al. 1977), is large and robust, but essentially modem in overall morphological configuration (Howell 1978; Kennedy 1984; Stringer 1986). However , the right innominate has an acetabulocristal buttress of cortical bone , which is present on a number of Lower and Middle Pleistocene hominid innominates from Africa and Europe (Stringer 1986). The dating of the Kabwe material is problematical because of the uncertainty as to the contemporaneity of the individual hominid remains with each other , and with the fauna} material and the terminal Acheulean stone artefacts. Stringer ( 1986: 115) suggested that only the left tibia (E.M. 691 ), and a femoral shaft fragment (E.M. 793) appear to have been found in close proximity to Kabwe 1, and that the remainde r of the material was "recovered in various circumstance s and must be considered unprovenanc ed". The cranium, along with other material, was recorded as having been recovered from a pocket of lead carbonate ore, yet assays showed that the skull and a tibia fragment contained more zinc than lead while the other bones had a predominance of lead, and man; of the stone artefacts were incrusted with lead ore (Clark et al. 1950). This may have been due to local variations in the mineral constituents of the soil. Fluorine and nitrogen analyses suggest that the bone material is of the same age (Oakley et al. 1977).

Kabwe-Broken Hill-Rhodesian Man Stone artefacts, fauna} material and hominid remains were recovered during mining operation s at a lead/zinc mine at Broken Hill, Northern Rhodesia (now Kabwe , Zambia , Fig. 2: 1; Hrdlicka 1930). The hominid remains consist of an almost complete cranium , fragment of a right maxilla, a parietal fragment and a number of postcranial bones. Kabwe 1, an adult cranium (Plate 1; Drennan 1953, 1955; Howell 1978; Wolpoff 1980a), is very robust with a low vault, marked postorbital constriction, and mid-facial and alveolar prognathism. The supraorbital torus is massive, and continues laterally forming a robust frontal process . The receding frontal squama is markedly keeled, while the short and relatively flat parietals display what may be called angular tori, postbregmatic and pre-lambdoid depressions, little bossing , and a high degree of mid-sagittal keeling. The greatest cranial breadth is found in the mastoid region , and the sides of the vault are flattened. The occipital is strongly flexed with a prominent , continuous and sharply defined torus that overhangs the flat and almost horizontal nuchal region. The cranial base is essentially modem in that it has a short basioccipital and steep clivus (Laitrnan et al. 1979).

Out of twenty four mammal species in the fauna! sample six are extinct which indicates a pre-Upper Pleistocene date (Klein 1973). Vrba (1982) has suggested that the bovid remains from Kabwe suggest a date slightly younger than Saldanha, whereas Partridge ( 1982) has proposed that the cranium is probably older than either the faunal assemblage or the bulk of the stone artefacts. An amino acid racemisation date of 110,000 years BP was obtained for a femoral fragment from Kabwe (Bada et al. 1974 ), but must remain questionable because the racemisation rate was calibrated with radiocarbon dated bone from Mumbai Cave, 150km way (Fleming 1976). McBrearty and Brooks (2000) cite preferred age estimates of 780,000- 1.33 m.y.a. and 1.07- 1.33 m.y.a. for the Kabwe hominids based on associated fauna .

The face is large and robust with a wide interorbital area moderately sized nasal aperture, large rectangular orbits, and an expanded maxillary region without canine fossae. The mid-facial region displays a relatively marked degree of prognathism (Stringer 1978). The zygomatics shelve relatively evenly into the maxillae , but there is an area of swelling behind the zygomaxillary suture (Rightmire 1996). Th~ me~ial section of the frontonasal suture is relatively honzontal. The palate is deep with steep walls in the molar region, and a great distance between the molar rows. The maxillary teeth are heavily worn and display deep caries. It has been suggested that the Kabwe 1 individual may have suffered from lead poisoning, something that is often associated with marked caries (A. Bartsiokas pers . comm.) .

The fauna} material, artefact assemblage and the morphology of Kabwe 1 suggest a date of possibly 200-400,000 years BP for at least some of the hominid material. Being so complete, Kabwe 1 was one of the core group of fossils that made up the basic data set that was used in all of the cranial metric analyses conducted (Table 1:2). For this reason, the exact placement of it varied somewhat depending on the hominids included in the analysis . When no subSaharan African fossils were included Kabwe 1 grouped with archaic Homo sapiens from Europ e such as Petralona , Arago 21 and Steinheim, or material from western Asia and north Africa such as Skhiil 5 and Jebel Irhoud 1 (for example Figs . 4:3, 4 :7, 4:8, 4:9, 4 : 10, 5:7, 5:8, 5:9, 5: 10, Tables 4 :5, 5:3, 5:4). When sub-Saharan African fossils were added to the analyses Kabwe 1 generally clustered with them. These included Bodo 1, Saldanha 1, Ndutu , Eliye Springs 11693, Florisbad, Singa, Laetoli H18 and Omo 2 (Figs. 2:3, 2:4, 2:9,

~abwe 2 (Howell 1978; Stringer 1978, 1986; Wells 1947), a nght maxilla with damaged second molar and a third molar preserved , is similar to Kabwe 1 in a number of respects including the external palate breadth. However , it is smaller with a shorter subnasal region , a shallower palate , more transverse orientation of the zygomatic process, a higher zygomaxillare angle, a larger nasal breadth, a larger third molar, and a distinct, but shallo w, canine fossa. These differences may be due to sexual dimorph ism, with Kabwe 1

30

Kabwe 2 is similar to Laetoli H 18 in canine fossa morphology, subnasal dimensions, and degree of prognathisrn, but it is larger in palate breadth, na~al brea~th, and in third molar size (Stringer 1986). The shght camne fossa development on Kabwe 2 also resembles Ndutu, Florisbad, Eliye Springs 11693 and Steinheim. Brauer (1984a) placed Kabwe 2 with Omo 2, Laetoli J:Il8 and Florisbad in his Homo sapiens Grade 2 (late archaic Homo sapiens).

Tables 2:2, 2:3, 2:4, 2:5). On the occipital analysis (Fig. 5:14) Kabwe 1 was positioned somewhat away from the other archaic hominids. It should be noted, however, that most of the measurements for Kabwe 1 used in this analysis were estimates, which could be affecting its placement. In earlier analyses Kabwe 1 grouped with Skhiil 4 and 9, Petralona, Zuttiyeh, Saldanha 1 Jebel Irhoud 1 and 2 (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). On the analysis of non-metric traits Kabwe 1 occupied an individual position and was not grouped with other hominids. Closest to it was Steinheim, and slightly further away the "classic" Neanderthals from Europe (Table 1:3, Fig. 5:15). This analysis is consistent with the results of a study of derived Neanderthal features carried out by Santa Luca ( 1978), which also distinguished Kabwe 1 from the Neanderthals.

Eyasi A number of hominid remains were recovered from the surface near the northeastern end of Lake Eyasi, Olduvai Gorge in Tanzania (Fig. 2:1; Mehlman 1984, 1987; Protsch 1976b, 1981). Eyasi 1 is a fragmentary calotte (Protsch 1976b, 1981). It has marked postorbital constriction, no sagittal keeling, a relatively low flat frontal and a slightly flexed occipital squama. The supraorbital region is moderately developed and the occipital torus is only prominent centrally. Eyasi 2 is an occipital fragment with greater flexion of the occipital than Eyasi 1 and a more prominent torus (Protsch 1981). Eyasi 3 is also an occipital fragment, but provides little morphological information (Protsch 1981).

These results are comparable to those of Corruccini (1974a) and Stringer (1974a, 1978). Van Vark (1983a, 1983b) found Kabwe 1 to group with Petralona, Steinheim and Arago 21. Brauer (1984a) found that on his principal components analysis of the frontal squama Kabwe 1 fell near Saldanha 1 and Laetoli Hl8, whereas on his analyses of the parietals it fell by itself, although Saldanha 1 was not too far removed. On the analysis of the splanchnocranium Brauer (1984a) found that Kabwe 1 was well separated from Petralona and Bodo 1 on the second axis. Howells ( 1989) found Kabwe 1 to be grouped with J ebel Irhoud I and Skhiil 5 and separated from "classic" Neanderthals from Europe.

Because these remains were surface finds their dating is difficult to establish. Aspartic acid racemisation dates of 34,000 years BP and 35,000 years BP have been proposed for Eyasi 1 and 35,000 years BP for Eyasi 2 (Protsch 1975, 1976a, 1976b, 1981), with Mehlman (1987) providing calibrated amino acid racemisation dates of 35,600 years hp for Eyasi 1, and 34,000 years bp for Eyasi 2. There are, however, some major problems surrounding these dates (Mehlman 1984, 1987). Bones which have been burnt, exposed to intermittent leaching by water and contain little nitrogen like the Eyasi hominids can produce erroneous, often inaccurately young, aspartic acid dates (Hare 1980; Mehlman 984, 1987; Oakley et al. 1977; Protsch 1976b). As Mehlman has stated ...the Eyasi I fossils meet several criteria for achieving spuriously young racemization dates. (1987:141-2) Attempts at uranium series analyses of Eyasi 1 have been unsuccessful (Mehlman 1987).

The results are consistent with the morphological pattern of Kabwe 1 in that it is similar to other sub-Saharan African hominids, especially Bodo 1 and Saldanha 1. These similarities have already been outlined. Other parallels are found for Kabwe on the Omo 2 calvaria, with similarities in the cranial contour, presence of mid-sagittal keeling, extensive nuchal regions, robust mastoid regions and in the marked occipital torus. Eyasi 1 also displays some resemblance to Kabwe 1 in cranial shape. Facially, Kabwe 1 is similar, although more robust, to Eliye Springs 11693 and Florisbad. Kabwe 1 is also similar to Petralona and Arago 21 from Europe in robustness and basic morphological configuration. The similarities with Petralona are quite marked, especially in the degrees of postorbital constriction and facial prognathism, the contour of the cranial vault, the degree of mid-sagittal keeling, occipital angulation and flexion of the cranial base, supraorbital torus development and overall facial form. Kabwe 1 is larger and more robust than Steinheim, which may be due to sexual dimorphism with Kabwe 1 being a male and Steinheim a female, but they do share archaic morphological features such as low cranial vaults, marked postorbital constriction, facial prognathism and large supraorbital tori.

Mehlman (1984, 1987) has conducted research at Lake Eyasi concluding that The stratigraphic sequence and taphonomic considerations indicate a middle Eyasi Beds provenience for the Eyasi I cranium and other W estbucht fossils. (1987:157-8) Mehlman (1987) felt that the youngest Eyasi Beds are reasonably dated to greater than 130,000 years bp, with the middle of the formation being 200,000 years old. He also identified at least seven extinct large mammals in the fauna from the Eyasi Beds, and found the artefacts to be unspecialised Middle Stone Age types with some Sangoan features. McBrearty and Brooks (2000) cite an age estimate of > 130 years BP for the Eyasi hominids based on extrapolated dates from overlying and underlying deposits and faunal correlations. The Eyasi hominids are probably of

Stringer et al. ( 1979) placed Kabwe, along with Bodo 1, in their Homo sapiens Grade 1. Brauer (1984a) placed Kabwe 1, along with Saldanha 1, Ndutu, Bodo 1 and Eyasi, in his Homo sapiens Grade 1 (early archaic Homo sapiens). McBrearty and Brooks (2000) placed Kabwe, along with Bodo, Ndutu, Saldanha and Homo erectus material into their Group 1.

31

upper Middle Pleistocene age. It is of interest that the nearby site of Mumba Rock Shelter has produced dental remains from Middle Stone Age deposits dated to between 130,000 and 110,000 years BP (Brauer 1984a). The fragmentary Eyasi 1 calotte was not included in any analyses . It is morphologically similar to the early robust hominids , Saldanha 1, Ndutu and Kabwe 1 in overall cranial form, and in some aspects Eliye Springs 11693, Laetoli H18 and to a lesser extent Omo 2. Brauer (1984a) placed Eyasi, along with Bodo 1, Kabwe 1, Saldanha 1 and Ndutu , in his Homo sapiens Grade 1 (early archaic Homo sapiens) . McBrearty and Brooks (2000) placed Eyasi, along with Bodo, Ndutu , Saldanha , Kabwe and Homo erectus material into their Group 1.

Omo 3 consists of a glabellar fragment and a frontoparietal vault fragment that are, where comparable, similar to Omo 1 (Day 1969, 1972). The Omo 1 remains were found partly in situ in Member 1 of the Kibish Formation in association with fauna and undiagnostic stone artefacts. The level above that which produced the hominid remains has been dated to 130,000 years BP by the uranium/thorium method using molluscs (Butzer et al. 1969). There are, however, problems with this method when used on molluscs in that less than 50% accuracy can be expected (Kaufman et al. 1971). The faunal material cannot be dated precisely , but suggests an upper Middle or early Late Pleistocene age (Day and Stringer 1982). Wolpoff (1980a), however , incorrectly states that the fauna suggested a date of 60,000 years BP . Omo 2 was a surface find some 2.5km from the KHS site of Omo 1. The sedimentary sequence at the KHS site and at the PHS site, where Omo 2 was found, are said to be the same (Butzer 1969). Butzer stated that ...it is not likely that the bones [of Omo 2] have been exposed for long, or that they have been lowered appreciably through erosion of the underlying sediments ... (1969:134) Butzer concluded that the two hominids, Omo 1 and 2, originally came from the same level - a minor disconformity in the upper third of Member 1 of the Kibish Formation (Butzer 1969). McBrearty and Brooks (2000) cite a preferred age estimate of > 130,000 years BP for Omo 1 and 2 based on the above uranium/thorium date.

Omo (Kibish) The remain s of three hominid s were recovered from near the Omo River in Ethiopia (Fig. 2: 1; Day 1969). Omo 1 consists of a partial skeleton with postcranial , cranial and mandibular remains (Plate 16). The incomplete vault includes parts of the occipital , parietals , frontal and temporal bones (Day 1969, 1972; Day and Stringer 1982). There are also facial and mandibular fragments . The frontal fragments, which do not join the parietals, suggest a long, and slightly curved morphology. The degree of postorbital constriction is moderate. The supraorbital region is not massively developed, whereas the glabella is quite prominent above the depressed nasal root. The supraorbital trigone would have been clearly definable from the superciliary arch. The parietals display a certain amount of bossing, and there is no mid-sagittal keeling or parasagittal flattening. The occipital is well-rounded with a moderate torus that fades laterally . The opisthocranion is located above the inion, indicating that the nuchal plane is reduced in size compared to the upper scale of the occiput. The mastoid processes are prominent and down-turned. The face is very fragmentary but appears to have been relatively robust, with some degree of prognathism . The maximum cranial breadth is located low on the parietals. The mandibular fragments include the symphyseal region , portions of the left corpus, and the right ramus with condyle . It is basically modem in size and robustness , with a well -developed mental eminence (Brauer 1984a; Day 1969, 1972). The fragmentary postcranial material is robust , but within the modem range of variation (Day 1969, 1972; Kennedy 1984).

The contemporaneity of these two fossils is still open to doubt, as is the 130,000 years BP date. Butzer (1989) reaffirmed the view that the dating of the Omo material was essentially correct, and suggested that Omo 1 and 2 belong to the early part of oxygen isotope stage 5, which Butzer (1982) dated to between 125,000 and 130,000 years BP (Table 5:1). Brauer (1989) contends that the suggested early dating of Omo 1 is supported by the amount of deposit between its find spot towards the base of Member 1 of the Kibish Formation and the radiocarbon date from the upper part of Member 3. This date of greater than 37,000 years bp (L1203-A) for Member 3 indicates that the Omo 3 fragments, which are from this member , are beyond the traditional range of the radiocarbon method (Butzer 1969, 1976, 1989; Oakley et al. 1977). The dating of Omo 1 and 2 remains inconclusive. Omo 1 was found to cluster with other sub-Saharan African hominids such as Omo 2, Kabwe 1 and Saldanha 1 (Table 2:4). This highlights a regional identity within the material. Omo 1 however, is not especially similar to this material, but would seem to be even less similar to the other crania in the analyses. In earlier analyses Omo 1 was a member of the same group as Florisbad , Borde r Cave 1, Skhiil 5, Jebel Qafzeh 6, Middle Pleistocene crania from Europe such as Arago 21, Steinheim, and Swanscombe , and Neanderthal crania including Krapina C, Saccopastore 1 and Gibraltar 1 (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). Omo 1 is similar to Omo 2 in the supraorbital region where the superciliary arch can be differentiated from the supraorbital trigone, and in their slight degree of postorbital constriction. They differ in that Omo 1 has a more domed

Omo 2 is an almost complete calvaria (Plate 16; Day 1969, 1972). It is long and low, with the greatest cranial breadth on the mastoid region . Th e frontal is broad and flattened and there is some mid-sagittal keeling. Postorbital constriction is slight. The posterior root of the zygoma does not greatly overhang the auditory meatus . The occipital is angled and the opisthocranion coincide s with the inion. The upper scale of the occiput is curved , while the nuchal plane is flat. The prominent occipital torus has a flattened, sharply delineated surface. Laterally it joins marked supramastoid crests above prominent down-turned mastoid process es. The supraorbital region is moderately well-developed and the prominent superciliary arch can be separated from the more flattened supraorbital trigone.

32

vault, and has a less projecting occipital squama. The occipital of Omo 1 is also less robust and archaic than Omo 2 in that it has only slight muscle attachments on the reduced nuchal plane, a torus that fades laterally, and an expanded upper scale.

Omo 2 was found to cluster with Laetoli Hl8 (Fig. 2:5) and, on the k-means cluster analysis, with Kabwe 1 also (Table 2:4). On the correspondence analysis Omo 2 held a more individual position (Fig. 2:6). These groupings reflect high loadings on the variables that document the form of the frontal bone, especially its breadth (Fig. 2:6, Table A:2). When Laetoli Hl8 was not included in the analyses and more variables were used Omo 2 grouped with Petralona, Steinheim and Kabwe l. The results are compatible with those of Stringer ( 1974b, 1978), except he did not use Laetoli H 18, but are not in agreement with Brauer's ( 1984a, 1984b) parietal analysis where Omo 2 did not fall close to Laetoli H 18 or Kabwe 1. Omo 2 was not compared to the Ngandong crania or any of the earlier Homo erectus material from Africa or Asia, but in earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986) it grouped with the N gandong crania.

Stringer (1974b, 1978) found contrasts between the Omo crania. The generalised distance between the two crania was not great, suggesting that they do display certain similarities, which is consistent with the results of this investigation. In Stringer's analyses the affinities of the two Omo hominids, however, were quite different, with Omo 1 falling closer to anatomically modem crania such as Skhul 5, and Upper Palaeolithic crania from Europe, while Omo 2 was closer to more archaic crania such as Kabwe 1 and the N gandong material (Stringer 1974b, 1978). Day and Stringer ( 1982) compared Omo 2 and their new reconstruction of Omo 1 ( which was only marginally different in frontal dimensions and cranial length to the earlier reconstruction used by Stringer [1974b, 1978]), to their definitions (based on derived characteristics) of anatomically modem Homo sapiens, Homo erectus and Neanderthals. They found the two Omo crania to differ markedly in their affinities, with Omo 1 aligned with anatomically modem Homo sapiens, and Omo 2 with Homo erectus.

This result was consistent with the results of Stringer (1974b), Andrews and Williams (1973), and with Day's ( 1969, 1972) initial interpretations of Omo 2. In these earlier analyses (Habgood 1982, 1984a, 1984b: Habgood and Walker 1986) the Omo 2 and Ngandong group also contained the Petralona cranium. On a pairwise cluster analysis Corruccini (1992) found Omo 2 to group with Jebel Irhoud 1 and 2 Jebel Qafzeh 6, Sk.hiil4 and 9 and Florisbad.

On Brauer's ( 1984a, 1984b, 1984c) analysis of the frontal bone, Omo 1 was, although separated somewhat from them, similar to Florisbad, Singa, Jebel lrhoud 2 and terminal Pleistocene African crania (Omo 2 was not included). Omo 1 was well separated from Jebel Irhoud 1, Laetoli Hl8, Kabwe 1, Saldanha 1 and Border Cave 1 in this analysis (Brauer 1984a). On Brauer's (1984a) principal components analysis of the parietals Omo 1 was placed closest to Omo 2, Boskop and Jebel Irhoud 1. It was again well separated from Saldanha 1, Kabwe 1 and Laetoli Hl8 (Brauer 1984a). Brauer's (1984a) results are not inconsistent with those reached here. During the current study Florisbad, Singa and Boskop were not included in any analyses with either Omo 1 or Omo 2. On a principal components analysis using vault measurements, Brauer and Rimbach ( 1990:Figure 5) found Omo 1 to be separated from the other material in the analysis, with Cro-Magnon 1 closest to it. It is worth noting that Omo 1 was well separated from Eliye Springs 11693, Laetoli Hl8 and especially Sk.hiil5.

Table 2:4. K-means cluster analysis with Laetoli Hl8, Omo 2, Saccopastore 1 and Petralona. Analysis includes 77.7% of the total within-group variance. Group 1 Kabwe 1, Omo 2, Laetoli Hl8. Group 2 Predmost 3, Predmost 4, Cro-Magnon 1, Mladec Chancelade, Abri Pataud 1, Dolni Vestonice 3.

1,

Group 3 Steinheim, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Petralona, Saccopastore 1. Group 4 Sk.hiil5, Combe Capelle, Oberkassel 1, Oberkassel 2.

The morphological affinities of the two Omo crania are consistent with the above mentioned results in that they do have similarities with each other. Omo 1 has, generally, modem affinities, but in a number of features it is reminiscent of possibly contemporaneous archaic fossils. With Eliye Springs 11693 it shares relatively curved parietals, and a rounded occipital with a reduced nuchal plane and an expanded upper scale. Laetoli Hl8 is also similar to Omo 1 in having a rounded, although less protruding, occipital with a reduced torus, parietal bossing, and a moderately developed supraorbital region with superciliary arches separated from the supraorbital trigones. Brauer ( 1984a, 1984b) has suggested that Omo 1 displays similarities with the "Afalou type" from north Africa, and the "Cro-Magnoid type" from Europe, but does not outline these similarities in any detail.

Brose and Wolpoff ( 1971) found Omo 1 to be essentially modem in both metrical and morphological features, but then suggested that it was extremely similar to Amud 1, a Neanderthal cranium. Omo 1 and Amud 1 were not included in analyses together, but their respective affinities do not suggest a close morphological affinity between them. Wells ( 1972) and Brauer ( 1984b) have suggested that Omo 1 exhibits morphological similarities to anatomically modem fossils from north Africa and Europe. Such a relationship was not highlighted by this investigation in that Omo 1 clustered with archaic crania and not with fully anatomically modem ones. Their suggestion, however, is more compatible with the results of earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986) where Omo 1 grouped with skulls such as Sk.hiil 5, Jebel Qafzeh 6 and Border Cave 1, but these crania are still not fully anatomically modem.

Omo 2 is also quite similar to Laetoli H18. They both have low, flat frontals with superciliary arches separated from the

33

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Figure 2:6. Correspondence analysis with Omo 2 and Laetoli H18.

34

supraorbital trigones, comparable sagittal curvature of the parietals, and similar robust mastoid regions. However, the form of the occipital squama is different. In this region Omo 2 is closer to Saldanha 1 and Kabwe 1, in that it has an angled occipital with a prominent torus and an extensive nuchal region. The sharply defined occipital torus of Omo 2 is reminiscent of the more prominent torus found on Kabwe 1. The angle of the nuchal plane is not as horizontal as that of Kabwe 1, but is close to that of Saldanha 1. Omo 2 resembles Eliye Springs 11693 in the position of maximum cranial breadth, the presence of temporal crests, an extensive nuchal plane and a reduced occiput, robust mastoid region, the limited overhang of the auditory meatus by the posterior root of the zygoma, the broad frontal, and slight postorbital constriction. There are also similarities between Omo 2 and Eyasi 1 in cranial contour, but in other aspects, such as occipital torus form, they are different. The sagittal ke~ling of Omo 2 is reminiscent of Kabwe 1 and Bodo 1, but differs from Saldanha 1, Ndutu, Eyasi 1 and Laetoli H18 that do not have this feature. The broad frontal of Omo 2 is comparable to that of Florisbad, and differs from the anteriorly constricted frontals of Bodo 1, Kabwe 1, Saldanha 1, Ndutu, and Eyasi 1. This combination of archaic and more advanced features may be the reason for Omo 2 falling on its own on the correspondence analysis (Fig. 2:6). Thoma (1978a) has suggested that Omo 2 may represent the ancestral type froi:11 which anatomically modem forms developed. This relationship has not been supported in that Omo 2 clusters with archaic forms. It is obvious that there are marked differences in the morphologies and affinities of Omo 1 and 2, which leads one to agree with Day and Stringer ( 1982) who thought the two differed more than one would expect if they were from a single population. This casts further doubt on the contemporaneity of the two individuals. Morphologically, one would expect Omo 1 to be more recent. An upper Middle Pleistocene date for Omo 2 would be consistent with its morphological affinities as it displays a mosaic _of morphological features that link in with both the earl~er hominids (Bodo 1, Saldanha 1 and Kabwe 1) as well as with the contemporaneous or possibly later hominids (Laetoli H18, Eliye Springs 11693 and Florisbad).

Floris bad An incomplete hominid cranium was recovered from the debris of an ancient spring on the southern slope of the Hagenstad salt-pan in the Orange Free State, South Africa (Fig. 2:1; Drennan 1935). The hominid remains consist of a frontal bone, parts of both parietals, portions of the right side of the face, and an isolated right upper third molar that probably belonged to the same adult individual as the other material. The reconstruction of the facial skeleton by Dreyer (1935) was inaccurate in that he joined all of the fragments together and then attached them to the frontal (Plate 3). This made the face very small, positioned the zygomatic fragment in an unnatural position producing a pronounced infraorbital hollow and caused the nasal cavity and the palate to be too narrow' (Clarke 1985). This means that many of the earlier interpretations of, and measurements from, the Florisbad cranium may be unreliable. A revised reconstruction of the Florisbad face has been produced (Clarke 1985) on which none of the fragments join, and the entire face is separated from the frontal (contra Dreyer [1935], and also Rightmire [1978a], who joined the nasal bones to the frontal). This separation has meant that most measurements of the face are problematical because the height and breadth of the orbit and the nasal cavity, the size of the interorbital region, and the position of the nasion can only be roughly estimated. Florisbad (Clarke 1985; Dreyer 1935; Galloway 1937; Rightmire 1978a; Wells 1972) has a broad and thick frontal bone with a relatively steep supraglabellar segment behind which it recedes gently. There is only slight postorbital constriction, and no indication of parietal bossing. There is a metopic ridge and a wide interparietal groove. The glabella is quite prominent and the supraorbital margin is thick and rounded, but does not form a complete torus in that the superciliary arch and the more flattened supraorbital trigone can be separated. There is no ophryonic groove, only a supraglabellar depression. The fragmentary face is slightly prognathic, and appears to have been quite broad with large biorbital and interorbital breadths. The preserved nasal bones are relatively flat and the nasion is not depressed. The present reconstruction (Clarke 1985) has only a small nasal aperture. The infraorbital region exhibits a slight canine fossa.

Stringer et al. (1979) place Omo 1, along with the material from Skhul and Qafzeh, in their Homo sapiens Grade 3b, whereas they place Omo 2 with Krapina, Gibraltar 1, Zuttiyeh and Florisbad in their Homo sapiens Grade 2 or 3. Brauer (1984a) placed Omo 1, along with the material from Klasies River Mouth and Border Cave, in his Homo sapiens Grade 3 ([early] anatomically modem Homo sapiens ), while he placed Omo 2 with Laetoli H 18, Florisbad and Kabwe 2 in his Homo sapiens Grade 2 (late archaic Homo sapiens). McBrearty and Brooks (2000) placed Omo 2, along with Florisbad, Laetoli H 18 and the Jebel Irhoud material into their Group 2. Omo 1, along with the material from Klasies River Mouth and Border Cave, was placed in their Group 3.

Wells (1972) suggested that the very broad frontal squama of Floris bad may be an indication of some malformation of the skull and that the specimen is physically abnormal. There are, however, no indications of any other abnormalities on Florisbad, and its broad frontal can be matched on other subSaharan African crania such as Omo 2. The dating of the Florisbad hominid is problematical because of the nature of the deposit it was recovered from, which consists of approximately seven metres of sand interleaved with four organic layers or peats, as well as pillars/cones of sand and debris produced by local spring-eye activity. As one spring-eye became choked with accumulated debris another would form. The fragmentary cranium, stone artefacts and fauna! remains were recovered from the debris cone of the western spring-eye at the level of Peat I (Dreyer 1938; contra Butzer [1984) who states that it came from the "eastern eye",

The Omo material is very important for an understanding of hominid evolution in sub-Saharan Africa, which makes it essential for the dating to be clarified (McBrearty and Brooks 2000).

35

Peats II and III. Beaumont (et al. 1978), however , suggested that the tools associated with the hominid remains, and others from Peat I, are Late Acheulean (Fauresmith) , which suggests a date greater than 170,000 years BP . The dolerite artefacts with sporadic flaking and bruising, which make up the so called "Florisbad Chopper Culture" (Meiring 1956), are probably natural objects which suffered abrasion and damage during periods of intense spring activity (Partridge 1982).

and Oakley [1954] and Coon [1962] who suggest that the skull lay in a depression in the top of Peat I). The western eye erupted through the overlying levels and into Peat III , and so the hominid could have come from any of these layers. Taphonomically, however , the hominid remains are consistent with the faunal material from Peat I. This fauna , including the hominid material , is taphonomically homogeneous in that it is generally large in size with limited fragmentation , whitish in colour , displays carnivore gnaw marks , and shows evidence of having weathered on the surface for some time prior to its incorporation into the deposit (J.S. Brink pers. comm.) .

As previously discussed , most splanchnocranial measurements obtained from Florisbad must be estimates because the facial fragments do not articulate with the frontal bone . Therefore any results obtained from analyses of metrical data are open to question . U sing estimated measurements , Florisbad was found to cluster, initially, with Kabwe 1 (the only other sub-Saharan African hominid in the analyses) , and then with Steinheim , Arago 1, Petralona , and Skhiil 5. The variables with high loadings document the form of the frontal squama , and the breadth of the interorbita l region and the nasal aperture . In earlier analyses Florisbad was a member of the same group as Omo 1, Border Cave 1, Skhiil 5, Jebel Qafzeh 6, Middle Pleistocene crania from Europe including Arago 21, Steinheim and Swanscombe, and Neanderthal crania such as Krapina C, Saccopastore l and Gibraltar l (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). In a principal components analysis of the frontal squama Brauer (1984a) found Florisbad to be well separated from Saldanha 1, Kabwe 1, Laetoli Hl8 , Jebel Irhoud 1 and also Border Cave 1. Of the archaic hominids it was positioned closest to Omo 1, Singa and Jebel Irhoud 2 (Brauer 1984a). On a pairwise cluster analysis Corruccini (1992) found Florisbad to group with Jebel Irhoud 1 and 2 Jebel Qafzeh 6, Skhul 4 and 9 and Omo 2.

Although the faunal sampl e from Peat I includes nine extinct large mammal species it is essentially modem , and is more recent than the Elandsfontein assemblage (Partridge 1982). Studies of the old faunal sample suggest that it dates to the end of the last interglacial or the early last glacial, somewher e around 115,000 to 110,000 years BP (J.S. Brink pers . comm.) . On geological grounds, Butzer ( 1984) suggested a later Middl e Pleistocen e date, possibly in excess of 150,000 years BP . Protsch (1975, 1976a) has dated Peat I to approximately 39,000 years BP by radiocarbon (38,680+/-2000 years BP [UCLA-l 745B] , 38,550 +/-3800 years BP [UCLA-1745C]) and amino acid racemisation measurements (39,000 years BP [4-UCLA-LJ]) on wood and bone, but there are doubts surrounding the stratigraphic provenance of the samples used (Rightmire 1978a). Other radiocarbon tests on samples from Peat I have indicated an age in excess of the effective range of the technique (Vogel 1970). Clarke (1985) reports two radiocarbon dates of greater than 47,200 years BP and greater than 44,600 years BP (no laboratory numbers given) for an organic clay layer just below Peat II. A date of more than 43 ,700 years BP (Pta-3465) has been obtained for a Middle Stone Age living floor between Peats II and III, and a preliminary uranium series date in excess of 100,000 years BP has been reported for Peat I (Clarke 1985).

The results confirm the similarity of Florisbad to other hominids from sub-Saharan Africa , and highlight its nonNeanderthal morphology (contra Drennan 1935, 1937). These results are in agreement with Galloway ( 1937) who also differentiated Florisbad from the Neanderthals. On the new reconstruction of Florisbad the interorbital and biorbital breadths approximate those of the more massive face of Kabwe 1. Florisbad also shares a broad upper face with Bodo 1 and Eliye Springs 11693. As outlined earlier , Florisbad is similar to Omo 2 in having a broad frontal bone without a continuous supraorbital torus, and shares with Laetoli Hl8 and Eliye Springs 11693 a number of features of the anterior portion of the frontal squama and of the maxilla . Similarities in the frontal bone are also found on Border Cave 1. The development of the canine fossa on the maxilla of Floris bad is also reminiscent of that of Ndutu and Kabwe 2. Wolpoff (1980a) also highlighted similarities of the face, and the contour of the frontal between Florisbad and Jebel Irhoud 1.

More recent dating at the site indicates a significantly earlier age for the Florisbad hominid (Griin et al. 1996). Electron spin resonance (ESR) dating was undertaken on in situ faunal dental material and from the original spring collection. Age ranges from 100-300,000 years BP were obtained , which may reflect re-working of material by spring activity . An age of 121,000+/-6,000 years BP was also obtained for the Middle Stone occupation horizon (Griin et al. 1996). ESR dating has also been undertaken using the hominid third molar, which provided an age of 259,000+/-35,000 years BP for the Florisbad hominid (Griin et al. 1996). Optically stimulated luminescence dating was also undertaken on quarts from the sediments at the site, which provided dates ranging from 279,000+/-47,000 years BP towards the bottom of the deposit to 146,000+/-15 ,000 years BP towards the top of the deposit (Griin et al. 1996). McBrearty and Brooks (2000) cite a preferred age estimate of 260,000 years BP for Floris bad based on the ESR dating of the hominid tooth.

Stringer et al. (1979) placed Florisbad in their Homo sapiens Grade 2 or 3 with Omo 2, Singa and Zuttiyeh. Brauer (1984a) placed Florisbad , along with Omo 2, Laetoli H18 , Eliye Springs 11693 and Kabwe 2, in his Homo sapiens Grade 2 (late archaic Homo sapiens) . McBrearty and Brooks (2000) plac ed Florisbad , along with Omo 2, Laetoli H l8 and the Jebel Irhoud material into their Group 2.

The stone artefacts associated with the hominid remains , and found throu ghout the stratigraphic succession below Peat III, are of Middle Stone Age affinity (Partridge 1982). As mentioned before , Clarke ( 1985) has excavated a Middle Stone Age living floor adjacent to the springs and between

36

Whilst an age of 270-300,000 years BP for ER-3884 is compatible with the morphology of the supraorbital region, it is less consistent with the relatively thin boned, broad and high posterior cranial vault. The minimum age estimates of approximately 160,000 years BP would be more compatible with the overall morphology of these Beret hominids and would place them with Laetoli H18 and the Herto and Jebel Irhoud material.

Ileret Cranial remains (KNM-ER-3884) and a femur fragment (KNM-ER-999) have been recovered from near Beret, East Turkana, Kenya (Fig 2.1; Brauer et al. 1992; Brauer et al. 1997). KNM-ER-3884 consists of a posterior cranial vault, supraorbital region, maxilla with most of the teeth and additional cranial fragments from an adult. Brauer et al. ( 1992) provided a provisional description of the hominid remains. The posterior cranial vault, which displays some post-depositional deformation, is quite broad and high with thin vault bones. There are weakly developed supramastoid crests and some lambdoidal protrusion. The occipital angle is relatively high. The temporal squama are quite low. Styloid processes would have been present. The cranial capacity is estimated to be approximately 1,400cc (Brauer et al. 1992). The supraorbital region is well-developed, especially the glabella segment, forms a continuous and slightly arched torus across the orbits and displays some lateral thinning. There is a well-defmed supratoral sulcus separating the supraorbital torus from the frontal squama. The maxilla is broad, relatively short and would have had a certain degree of sub-nasal prognathism.

The Beret KNM-ER-3884 hominid displays a mosaic of archaic and robust features and more modem features (Brauer et al. 1992). Brauer et al. (1997) contend that KNMER-3884 represents an archaic Homo sapiens or a transitional specimen. McBrearty and Brooks (2000) placed Beret KNM-ER-3884, along with Omo 2, Laetoli H18, Florisbad and Jebel Irhoud into their Group 2. Overall, ER3884 appears to be a robust ( especially the supraorbital region) archaic Homo sapiens individual.

Berto Fragmentary hominid remains, fauna} material and stone artefacts have been recovered from Herto Bouri, Middle Awash, Ethiopia (Fig. 2:1; Clark et al. 2003; Stringer 2003; White et al. 2003). The hominid remains include a relatively complete adult male cranium (BOU-VP-16/1 ), vault fragments from two other adult individuals (BOU-VP-16/2, 16/43) and the fragmentary skull of an immature individual (BOU-VP-16/5).

KNM-ER-999 is described as having a robust shaft, but with morphological similarities to material from the Levantine sites of Skhul and Jebel Qafzeh (Brauer et al. 1997). The hominid is thought to have derived from undifferentiated deposits attributed to the Guomode Formation, which overlies (or can be subsumed into) the Chari Member of the Koobi Fora Formation, (Brauer et al. 1992:Figure 3; Brauer et al. 1997). The hominid remains were initially given an age range of between 100-500,000 years ago by Brauer et al. ( 1992), who also indicated that the material could be considerably younger as it was recovered from close to the base of the Pleistocene/Holocene Galana Boi Formation. Two cranial fragments from KNM-ER-3884 and part of the femur have been dated by non-destructive '}'-rayspectrometry (Brauer et al. 1997). The two fragments from KNM-ER-3884 yielded U-Th ages of 272,000 years BP (minimum of 159,000 years BP) and 279,000 years BP (minimum of 162,000 years BP), while the femur provided an older age of 301,000 years BP (minimum of 205,000 years BP) (Brauer et al. 1997). The three samples also provided U-Pa ages of >180,000 years BP (Brauer et al. 1997).

The adult male cranium, Herto 1 (BOU-VP-16/1) is relatively complete except for the left facial skeleton (White et al. 2003). Herto 1 (White et al. 2003) is a large and robust cranium with a long and relatively high vault, a prominent glabellar region and a slightly receding frontal. There are high and arched superior temporal margins and only slight angular tori. The posterior of the vault reveals large mastoid processes, some occipital protrusion, ossicles within the lambdoidal suture and a strongly flexed occipital with a prominent occipital torus. The preserved right supraorbital region is robust, especially medially, thins laterally and is arched over the squarish orbit. The splanchnocranium is broad and flat in the midface with moderate alveolar prognathism. The interorbital area is wide, the nasal bones tall and relatively narrow, while the nasal aperture is broad with a sharp sill. There is an incisura malaris and a robust malar tubercle (White et al. 2003). The cranial capacity is estimated at 1,450cm3 (White et al. 2003). The dentition of Herto 1 is large and heavily worn.

The Beret KNM-ER-3884 hominid displays clear morphological similarities with Laetoli H18, especially in the form of the supraorbital region and maxilla (Plate 2). The supraorbital region is also similar to that on Herto 1 and the Florisbad hominid (Plate 3). There are limited parallels with Herto 1 in the size and shape of the posterior vault, with Herto 1 being more robust. ER-3884 is less archaic and robust than the Bodo and Kabwe hominids, but displays some similarities with Ndutu in the supraorbital region and Omo (Kibish) 2 in the size and shape of the posterior vault. There also appears to be morphological parallels with the north African Jebel Irhoud material (Plate 3). Brauer et al. ( 1992) identified morphological and metrical similarities with robust anatomically modem north African material from Afalou-bou-Rhummel and Taforalt and late Upper Pleistocene African material such as Lukenya Hill.

Herto 2 (BOU-VP-16/2) consists of thick vault fragments from a large and robust adult male cranium, whereas BOUVP-16/43 is a parietal fragment from a smaller individual (White et al. 2003). BOU-VP-16/5 is the fragmentary skull of an immature individual aged possibly 6-7 years (White et al. 2003). The Herto hominid material is said to display cultural modifications indicative of mortuary practice and curation (Clark et al. 2003). Herto 1 has a number of cut marks on the anteroinferior comer of the right parietal and on the supramastoid crest of the right temporal, whilst the fragmentary Herto 2 has evidence of bone modification (cut and scrapping marks) on 15 of its 24 fragments. BOU-VP-

37

16/5 displays cut marks on the basicranium and the edges around the missing occipital region near the foramen magnum are smooth and polished.

Laetoli-Laetolil (Ngaloba Beds) A hominid cranium, Middle Stone Age type artefacts, and fauna were recovered from the Ngaloba Beds at Locality 2, Laetoli, northern Tanzania (Fig. 2:1; Day et al. 1980).

The stone artefacts from Herto Bouri have elements of both Acheulian and Middle Stone Age technologies (Clark et al. 2003). The hominids , fauna and artefacts were recovered from the Upper Herto Member and have been dated by stratigraphic, geochemical and radio isotopic (the argon method) means to between 160,000 and 154,000 years BP (Clark et al. 2003) . Whilst the dating seems to be relatively robust, the younger date is based on stratigraphic correlations over a few 100 metres, the hominid material , apart from Herto 1, was recovered from the surface and the nature of the stone tools may suggest some mixing of material.

The adult hominid cranium, Laetoli H18 (Plate 2; Day et al. 1980; Magori and Day 1983), consists of an almost complete vault, portions of the cranial base and a detached maxillofacial fragment. The vault is long and low with only slight postorbital constriction. The frontal is low, except for a relatively steep supraglabellar region, and flat, with only a slight degree of sagittal keeling, and a moderately developed supraorbital region. The interorbital area is relatively wide, while the glabella is not prominent , and the brow-ridges are only moderately thick. The superciliary arch and the supraorbital trigone can be distinguished . The maximum cranial breadth is positioned low down on the relatively large parietal s, which display mid-parietal bossing. The rounded occipital protrudes and the expanded upper squama is quite high. The opisthocranion lies well above the inion. The mound shaped occipital torus is prominent but fades laterally. It is undercut, but has no clear upper margin. The mastoid processes are small with well-marked supramastoid and occipitomastoid crests, and deep digastric fossae.

Herto 1 was not available for inclusion in any multivariate analyses for this study. White et al. (2003:Figure 4) found that on a plot of the first two components from a principal components analysis, Herto 1 adopted an individual position , but was placed between archaic hominids and Neanderthals , and fossil and modem Homo sapiens. When compared to modem male samples, Herto 1 did not display any close relationship with modem African material, but was closer to modem Australian material although still clearly distinct and more archaic (White et al. 2003) .

The maxillofacial fragment is not exceptionally robust. There are slightly developed canine fossae on the maxilla, and the nasal aperture is not especially broad. The inferiolateral zygomaticoalveolar margin is horizontal. The subnasal region is prognathic, and the palate is relatively broad and deep.

Herto 1 is less archaic than the Bodo, Saldanha and Kabwe hominids, although Stringer (2003) has suggested similarities in the angulation of the occipital and form of the occipital torus. Herto 1 is larger and more robust in the occipital and supraorbital regions than the archaic Homo sapiens material from the sub-Saharan African sites of Omo Kibish, Florisbad, Laetoli, Eliye Springs and Klasies River Mouth and the western Asian sites of Skhiil and Jebel Qafzeh. There are no resemblances to Neanderthal material (White et al. 2003). Whilst there are some morphological similarities with Omo (Kibish) 2, the vault of Herto 1 appears to be higher, the frontal less flat and the occipital more angulated. There are similarities in the form of the supraorbital region between Herto 1 and Laetoli HIS (Plate 2), but the latter appears to have a shorter and lower vault and a more rounded and less protruding occipital. There are also similarities between Herto 1 and Ileret ER-3884 in the form of the supraorbital region large, but differences in the posterior vault. The closest morphological parallels appear to be with the possibly 130,000 years old Jebel Irhoud hominids , especially Jebel Irhoud 1 (Plate 3), from north Africa. Jebel Irhoud 1 has a relatively steep frontal , broad interorbital region, prominent mastoid processes, some occipital protrusion and a prominent occipital torus . Jebel Irhoud 1 does appear to have relatively longer and flatter parietals , lower temporal squama and more pre-lambdoidal flattening than Herto 1. Whilst Jebel Irhoud 1 has a smaller supraorbital region, a shorter face and a taller nasal aperture, it shares with Herto 1 a broad and flat face with squarish orbits, a broad nasal aperture , marked incisura malaris and alveolar prognathism .

The alluvial Ngaloba Beds at Locality 2, Laetoli, contain a water-worked vitric trachytic tuff, which is below the beds that yielded the hominid material that is tentatively correlated with the marker tuff in the lower unit of the Ndutu Beds at Olduvai Gorge. This is the only trachytic tuff younger than Bed IV at Olduvai Gorge and its age is estimated at 120,000+/-30,000 years BP (Day et al. 1980; Leakey and Hay 1982). A giraffe vertebra from the same bed that yielded the hominid remains has been dated by the uranium-thorium method to 129,000+/-4,000 years BP (Th-230) and 108,000+/-30,000 years BP (Pa-231) (Brauer 1989a, 1989b; Day 1987; Mehlman 1986). Brauer (1989b) also quotes an amino acid (isoleucine epimerization) date from animal tooth enamel from the hominid level of 100,000 to 200,000 years BP. McBrearty and Brooks (2000) cite preferred age estimates of 200-490 ,000 for Laetoli H18 based on argonargon ratio and palaeomagnetism . On the SSA cluster analysis, Laetoli H18 clustered with Omo (Fig. 2:4), while on the k-means cluster analysis the group included Kabwe 1 (Table 2:4). On the correspondence analysis Laetoli H 18 was clearly separated from the other hominids (Fig. 2:6). The positioning of Laetoli H18 was strongly influenced by the form of its frontal bone because the variables that cover this region had high loadings (Fig. 2:5, Table A:2) .

White et al. (2003 :745) concluded that the Herto hominid s represented "a population that is on the verge of anatomical modernity but not yet fully modem " and taxonomically classified them as Homo sap iens idaltu. They were regarded as the descendants of the earlier Homo rhodesiensis (Bodo and Kabwe) and the ancestor of anatomically modem Homo sapiens (White et al. 2003).

Magori and Day ( 1983) carried out a canonical variate analysis in which they compared Laetoli HIS to mean values of various groups, including a sub-Saharan early Upper Pleistocene group which included Kabwe 1, Saldanha 1, Omo 1 and 2, Eyasi 1, Florisbad , and Singa (a rather

38

morphologically diverse group). Their results were not particularly informative due to the nature of the groups with which they chose to compare Laetoli H 18, and because the analysis was dominated by overall size.

2 and Florisbad. McBrearty and Brooks (2000) placed Laetoli H18, along with Omo 2, Florisbad and the Jebel Irhoud material into their Group 2.

In a principal components analysis of the frontal squama carried out by Brauer (1984a), Laetoli H18 grouped with Saldanba 1 and Kabwe 1 and was well separated from Jebel Irhoud 2, Singa, Florisbad, Omo 1 (Omo 2 was not included) and Border Cave 1. In a principal components analysis of the parietals Laetoli H 18 was, although closest to Saldanba 1, separated from the archaic hominids including Kabwe 1, Omo 2 and Jebel Irhoud 1 (Brauer 1984a). On a principal components analysis of vault measurements, Brauer and Rimbach (1990:Figure 5) found Laetoli H18 to be separated from Eliye Springs 11693, Skhfil 5 and especially Omo 1, and with the immature Neanderthal Le Moustier 1 closest to it.

Eliye Springs An almost complete hominid cranium, KNM-ES 11693, was found near Eliye Springs on the western shore of Lake Turkana, Kenya (Fig. 2:1; Brauer and Leakey 1986a, 1986b). The supraorbital region is missing, as is much of the right temporal squama and the right half of the face, while the left half of the face is heavily damaged. There is also a certain degree of postmortem inward deformation on the right side of the cranium. The cranium (Brauer and Leakey 1986a, 1986b) is relatively long, low, and very broad. The maximum breadth is low down on the supramastoid crests. The vault bones of the matrix-filled cranium are quite thick. Postorbital constriction is slight. The frontal is relatively steep, especially the supraglabellar segment. There is a shallow ophryonic sulcus, and indications that the supraorbital region would have been similar to that seen on Laetoli Hl8. The parietals are relatively long and curved, with posterior flattening. Parietal bosses are quite well-developed. There is a marked heaping up of bone along the median plane on the frontal and continuing onto the parietals almost as far as the obelion depression. This heaping up of bone is extended more transversely than the mid-sagittal keeling, as seen on crania such as Kabwe 1.

The results of the cluster analyses are consistent with the morphological affinities of Laetoli H18, and highlight its resemblance to other sub-Saharan African hominids. Laetoli H18 displays a mixture of archaic (a low, flat frontal, supraorbital development, and robust mastoid region), and more advanced features (parietal bossing, a rounded occipital, reduced nuchal plane, and the separation of the superciliary arch and the supraorbital trigone ). This combination may account for the individual position held by Laetoli H18 on the correspondence analysis (Fig. 2:5). As previously outlined, Laetoli H 18 and Omo 2 are morphologically very similar, except for the occipital squama, where Laetoli H 18 is more rounded and less robust. Here, except for its protrusion, the Laetoli Hl8 occipital more closely resembles that of Omo 1 and Eliye Springs 11693, which also have reduced nuchal planes and expanded upper scales. Laetoli H 18 also resembles Eliye Springs 11693 in having slight postorbital constriction, temporal crests, a steep supraglabellar segment, well-curved frontal and parietals, and a slight canine fossa. The parietal bossing and form of the supraorbital region of Laetoli 18 are also reminiscent of Omo 1. Laetoli H 18 and Florisbad are similar in having little postorbital constriction, a steep supraglabellar region, lack of a continuous supraorbital torus, and a slight canine fossa. The steep supraglabellar region and parietal bossing of Laetoli H18 are also found on Border Cave 1. Jebel Irhoud 1, Maha, Zuttiyeh, Tabun Cl, and Skhfil 4 and 9 have similar supraglabellar regions. Apart from the steeper supraglabellar region and the protrusion of the occipital, the sagittal contour of Laetoli H 18 is similar to that of Eyasi 1. The development of the canine fossa on Laetoli H 18 is reminiscent ofKabwe 2, and to a lesser extent Ndutu.

In norma occipitalis the parietals gently taper until they reach prominent temporal crests, where they sharply converge towards the mid-sagittal plane. The occipital squama displays an expanded and strongly curved upper scale and a horizontal nuchal plane. There is a certain degree of lambdoidal flattening and only slight occipital protrusion. There is no occipital torus, but there is an oval depression corresponding to the suprainiac fossa. The damaged mastoids would have been relatively short and broad, with prominent occipitomastoid and supramastoid crests. The posterior root of the right zygoma does not greatly overhang the auditory meatus. The minimum frontal breadth is located relatively far back on the temporal crests. The face would have been very broad and quite short, with a certain degree of prognathism. The rectangular orbits are relatively large. The nasal bones are small and form a distinct angle to each other giving a pinched appearance, while the nasal aperture is low and rounded. The right maxilla has a shallow canine fossa.

Morphologically (and chronologically?) Laetoli Hl8 would appear to be intermediate between the robust and archaic crania (Bodo 1, Saldanha 1, Ndutu, and Kabwe 1), and the more advanced crania (Omo 1 and the material from Border Cave and Klasies River Mouth) from sub-Saharan Africa. A similar intermediate position is held by Florisbad, Omo 2, Eyasi 1, Eliye Springs 11693, Ileret and the Herto hominids. Rightmire ( 1989) has suggested that the primitive .feature~ of Laetoli H 18 have been over stated and that 1t provides evidence for a modern morphological presence in east Africa. Brauer (1984a) placed Laetoli Hl8 in his Homo sapiens Grade 2 (late archaic Homo sapiens) along with Omo

The cranium, along with faunal material, which does not include any extinct species, was found in reworked beach deposits, and so no date for it is available (Brauer and Leakey 1986a, 1986b). The heavy mineralization of the Eliye Springs 11693 has been interpreted as indicating that it derived from deposits predating the Holocene beds (Brauer and Leakey 1986a, 1986b). Brauer et al. ( 1997) gave Eliye Springs 11693 a provisional age of around 200,000 years BP, however, all that can be said at present is that the specimen was most probably derived from Upper Pleistocene deposits.

39

H 18 and Omo 1. A reduced nuchal plane and an expanded occiput can also be found on Ndutu , Laetoli Hl 8 and Omo 1. The broad face of Eliye Springs 11693 is matched by Bodo 1, Kabwe 1 and Florisbad , while a shallow canine fossa can be found on Ndutu, Kabwe 2, Laetoli Hl8 and Florisbad. The lack of overhang of the auditory meatus by the posterior root of the zygoma is similar to the pattern found on Omo 2, and differs from Kabwe 1 (Brauer and Leakey 1986a, 1986b). It is also of interest to note that the overall cranial contour of Eliye Springs 11693 is reminiscent of the Dali cranium from China.

Eliye Springs 11693 was found to cluster with Saccopastore 1 and Kabwe 1 (the only other sub-Saharan African hominid in the analyses) and Petralona and Steinheim (Fig. 2:7, 2:8, Table 2:5). It was well separated from the Neanderthals and the anatomically modem crania . There were high loadings on the variables that document facial size, cranial height, midsagittal curvature, asterionic breadth and temporal height (Fig. 2:8, Table A:3). Table 2:5 . K-means cluster analysis with Eliye Springs KNM-ES 11693. Analysis includes 78.7% of the total within-group variance.

There are, however, a number of features found on Eliye Springs 11693, such as the heaping up of bone along the median plane , presenc e of a suprainiac fossa, and the great cranial breadth , that are not matched by any of the other subSaharan African hominids . Computer tomography of Eliye Springs 11693 has revealed thickening of the vault, especially the diploe component, reaching up to 20mm at the posterior parietals . Changes to the vault may have been caused by a pathological condition such as severe chronic anaemia (Brauer et al. 2003). The symmetrical enlargement of the frontal and parietal bones, which does not match sagittal keeling, and the porotic nature of the vault surface are consistent with this assessment. It is difficult to determine if this pathological condition has had any significant impact on the placement of this individual on the various multivariate analyses.

Group 1 Skhfil 5, Predmost 3, Cro-Magnon 1, Mladec 1. Group 2 Steinheim, Kabwe 1, Petralona , Saccopastore 1, Eliye Springs KNM-S 11693. Group 3 Gibraltar 1, La Chapelle 1, La F errassie 1.

Brauer (Brauer and Leakey 1986a, 1986b) included Eliye Springs 11693 in a number of limited principal components analyses which dealt with individual portions of the cranium, comparing it to archaic and modem crania (mostly from the north African sites of Afalou and Taforalt) from, predominantly, Africa. On the analysis of the face Eliye Springs 11693 fell near Petralona and contrasted with Border Cave 1 (the only other non-modem hominid in the analysis) and fully modem crania from north Africa. In the analysis of the frontal squama, Eliye Springs 11693 was placed in an intermediate position between the archaic crania (Petralona , Kabwe 1 and Laetoli H18) , and most of the fully modem crania. On both the parietal and occipital analyses Eliye Springs 11693 fell with archaic hominids and contrasted with the fully modem crania. These analyses, however , appear to be dominated by overall size. On a principal components analysis of vault measurements , Brauer and Rimbach ( 1990:Figure 5) found Eli ye Springs 11693 to be separated from Laetoli H18, Skhfil 5 and Omo 1, and with late Pleistocene north African material closest to it.

Brauer (Brauer and Leakey 1986b) placed Eliye Springs 11693 in his Homo sapiens Grade 2 (late archaic Homo sapiens).

Singa A calvaria was found eroding from a deposit within the "Gezira clay" exposed in the west bank of the Blue Nile in the Singa district of eastern Sudan (Fig. 2: 1; Oakley et al. 1977; Stringer 1979). The calvaria is short, quite low, relatively broad at the base, has only a moderate degree of postorbital constriction, and has a "heart-shape" when viewed in norma occipitalis and norma verticalis. Almost 80% of the vault thickness is made up of diploe (Webb 1989). The frontal is steep in the supraglabellar region, but then gently recedes towards the bregma . The supraorbital region is moderately developed. The superciliary arches are medially separated from each other and laterally from the thinner supraorbital trigones. The parietals are short and flat with a high degree of bossing, while the protruding occipital is rounded in profile . The upper squama of the occiput is longer than the poorly defined nuchal plane. The greatest cranial breadth is high up on the parietals .

The results from the various multivariate analyses are generally consistent with the overall morphological configuration of Eliye Springs 11693 and confirm its archaic nature. They do not, however , highlight any strong similarities between Eliye Springs 11693 and Laetoli H18. Eliye Springs 11693 displays a mosaic of cranial features , but has most similarities with Omo 2 and Laetoli H 18 (Brauer 1989; Brauer and Leakey 1986a, 1986b). A low position of maximum cranial breadth is also seen on Ndutu , Kabwe 1 and Omo 2, as is a robust mastoid region . Short, broad mastoid processes are also present on Omo 2 and Border Cave 1. Slight postorbital constriction is also found on Omo 1 and 2, Laetoli H18 , Florisbad and Border Cave 1, while prominent temporal crests occur on Kabwe and Omo 2 and, to a lesser extent, Laetoli H 18. The mid-sagittal contour of the frontal of Eliye Springs 11693, especially the steep supraglabellar segment, is similar to that of Laetoli Hl 8 and Florisbad . There is also an overall resemblance to the frontal of Saldanha 1. Parietals curved to a similar degree as on Eliye Springs 11693 are found on Laetoli H18 and Omo 1, while the occipital contour is reminiscent of Eyasi 1, Laetoli

A precise date for the Singa calvaria is uncertain. There are possible contamination problems with a radiocarbon date of 17,300+/-200 years BP (in Stringer [1979] this date is 17,300+/-2000 years BP) obtained from a crocodile tooth from a corresponding level at the nearby site of Abu Hugar (Oakley et al. 1977; Stringer 1979). Relative dating indicates that the Singa calvaria is contemporaneou s with the Upper Pleistocene fauna from Abu Hugar (Oakley et al 1977). The stone artefacts from Singa and Abu Hugar had been identified as a Middle Stone Age (Proto-Stillbay) industry (Oakley et al. 1977), but have also been related to the Final

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41

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Acheulean (Brauer 1984a). Marks (1968), however, has suggested that the artefacts are nondescript , and although stratigraphically equivalent to the calvaria, they may not be of the same age.

Table 2:6 . K-means cluster analysis with Singa. Analysis includes 71.4% of the total within-group variance .

Group 1 Mladec 1, Dolni Vestonice 3, Oberkassel 1, Oberkassel 2, Abri Pataud 1.

Griin and Stringer ( 1991) attempted to obtain an electron spin resonance (ESR) age for the Singa hominid . They analysed two mammalian teeth from the site that were found to contain high amounts of uranium and so produced significantly different ages for early uptake (EU) and linear uptake (LU). Provisional dates of 97,000+/-15 ,000 years BP (EU) and 160,000+/-27,000 years BP (LU) were obtained (Griin and Stringer 1991).

Group 2 Gibraltar 1, La Chapelle 1, La Ferrassie 1. Group 3 Predmost 3, Predmost 4, Combe Capelle, Cro-Magnon 1, Chance lade.

Calcrete matrix adhering to the external surface of the calvaria and enclosing associated mammal teeth have been dated by mass-spectrometric uranium-thorium (U-Th) (McDermott et al. 1996). The associated dental matrix provided dates ranging betw een 129,000+/-5,000 years BP and 143,000+/-27,000 years BP, whereas the calvaria matrix samples provided dates of 133,000+/-26,000 years BP and 145,500+/-7500 years BP (McDermott et al. 1996). An age of 133,000+/-2,000 years BP was obtained from a sample that was essentially uncontaminated by silicate detritus and provides what is described as a "robust" U-Th age (McDermott et al. 1996). An Equus tooth produced massspectrometric U-Th ages of 72-90,000 years BP, and provided ESR dates of 85-97,000 years BP (EU) and 140159,000 years BP (LU) (McDermott et al. 1996). McDermott et al. ( 1996) concluded that their results provided an age of at least 133,000 years BP for the Singa hominid. McBrearty and Brooks (2000) cite a preferred age estimate of 190130,000 years BP for the Singa calvaria based on these Useries dates .

Group 4 Steinheirn, Kabwe 1, Skhiil 5, Singa . parietal angles and indices Stringer (1978) did not find Singa to be separated from the other Middle and early Upper Pleistocene hominids . The steep supraglabellar region of the frontal squama of Singa is similar to Laetoli H 18 and Florisbad , while a separation of the superciliary arch from the supraorbital trigone is found on Omo 1 and 2, Laetoli H 18 and Florisbad. A rounded occipital profile and parietal bossing can also be found on Omo 1 and Laetoli Hl8. Although there are some morphological similarities between the Singa calvaria and anatomically modem humans, it does present numerous archaic features such as the development of the supraorbital region, the form of the temporal bones, the meningeal vessel patterns and the small cranial capacity and endocranial proportions (Tobias 1968; Stringer 1979; Stringer et al. 1985). The posterior portion of the cranium , however, is unusual , and has prompted comparisons with the late Pleistocene Boskop calotte, and modem Khoisan crania (Briggs 1955; Wells 1972; Woodward 1938).

Singa was found to cluster with Steinheirn, Skhiil 5 and Kabwe (Fig. 2:9, Table 2:6), while on the correspondence analysis it fell on its own (Fig. 2: 10). When only a limited number of variables were used, Singa was a member of a group that included Steinheim and three European Neanderthal crania (Table 2:3). There were high loadings on the variables that document the form of the posterior portion of the cranium (Fig. 2: 10, Table A:4). The non-metric analysis placed Singa with the anatomically modem crania (Table 1:3, Fig. 5: 15).

Others have questioned these comparisons (Greene and Armelagos 1972; Tobias 1968; Rightmire 1975a), with Broth well ( 1974) suggesting that the unusual cranial shape might be the result of a longitudinal growth defect affecting the parietals. Brothwell's (1974) proposal is supported by the excessive diploic thickening at the parietal bosses and the marked development of the sphenoidal sinus of Singa, which suggest some form of pathological alteration, although a specific pathological disorder has been difficult to identify (Stringer et al. 1985). Webb (1989 , 1990) observed that Singa had a morphology that was similar to crania that have symmetrical osteoporosis , although there is no pitting on the outer table of bone that normally accompanies this condition. Webb (1989 , 1990) suggested that Singa suffered from some form of haemoglobinopathy or severe and chronic anaemia. The unusual morphological configuration may account for the divergent position of Singa on the correspondence analysis (Fig. 2:10) and also on Brauer's (1984a) parietal analysis (although this was not found by Stringer [1978]).

Stringer ( 1979), using Penrose size and shape statistics, found Singa to be closest to the Zhoukoudian Homo erectus and Ngandong samples , Jebel Irhoud 1, Saccopastore 1, Kabwe 1 and Omo 2. He concluded that its affinities lay with the Omo and Jebel Irhoud crania (Stringer 1979). Wells (1972) and Rightmire (1975a) have also suggested that Singa displays similarities with the Omo crania . In a principal components analysis of the frontal squama carried out by Brauer (1984a) , Singa was well separated from most of the other archaic hominids and was positioned near the origin of the plot with late Pleistocene hominids from Africa. Of the archaic hominids , J ebel Irhoud 2 was the nearest to Singa (Brauer 1984a). On a principal components analysis of the parietals Brauer ( 1984a) found Singa to be well separated from all other hominids . However, on plots of

Stringer et al. (1979) placed Singa in their Homo sapiens Grade 2 or 3 along with Omo 2 and Florisbad. Brauer (1984a) provisionally placed Singa with Omo 1 and the hominids from Border Cave and Klasies River Mouth in his

42

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43

grave along with a perforated Conus shell (an ornament or pendant?) , appears to have been covered in red ochre as some of the bones have reddish-brown stains (Beaumont 1980; Cook et al. 1945). Morphologically , the skeleton is essentially modem (de Villiers 1973).

Homo sapiens Grade 3 ([early] anatomically modem Homo sapiens ). McBrearty and Brooks (2000) placed Singa, along with Laetoli H18, Omo 2, Florisbad and the Jebel Irhoud material into their Group 2.

Whilst the Singa calvaria displays both archaic and more modem features its usefulness in an examination of hominid evolution in sub-Saharan Africa is limited because it may display a pathology affecting its cranial form.

Border Cave 5 consists of an almost complete adult mandible containing a number of teeth (Beaumont 1980; de Villiers 1976). The relatively short ramus is almost vertical and quite broad , while the corpus is of moderate height and robusticity. The symphyseal region displays a well-developed mental protuberance that is more prominent than the eminence on Border Cave 2.

Border Cave A number of hominid remains have been recovered from Border Cave, which is located just below the eastern scarp of the Lebombo Mountains , Ingwavuma District, KwaZulu , in northern South Africa (Fig. 2:1; Morris 1992; Wells 1950). The cave contains a long archaeological sequence from the Middle Stone Age (MSA) onwards. The remains of at least four hominids are said to have derived from the MSA levels at the site (Beaumont 1980; Beaumont et al. 1978; Cooke et al. 1945; Morris 1992). Border Cave 1 and 2 were extracted from the cave deposits by guano diggers , Border Cave 3 was recovered during systematic excavation at the site, and Border Cave 5 was found eroding out of a standing section (Border Cave 4 is of Iron Age date [Beaumont 1980]). Postcranial material was also reported from the site, but has been lost (Morris 1992). Additional postcranial material, including fragments from a right proximal ulna (BC7) , a right humeral shaft (BC6) and two right metatarsals (BC8a & b ), have been recovered from sediments that had slumped into earlier excavations (Morris 1992; Pearson and Grine 1996).

The remaining postcranial materia l consists of fragments from a robust right humeral shaft, a right proximal ulnar and two metatarsal (Morris 1992; Pearson and Grine 1996). As previously mentioned , Border Cave 1, 2, 3 and 5 are said to derive from Middl e Stone Age levels at the site (Beaumont 1980; Beaumont et al. 1978; Butzer et al. 1978; Cooke et al. 1945). However, this attribution is not as secure as is often suggested (for example Brauer 1984a, 1984b, 1984c; Rightmire 1984a). The method of recovery of Border Cave 1 and 2 (guano removal) means that their exact provenance will probably never be known. The character of soil adhering to interstices on Border Cave 1 is said to match a distinctive chocolate coloured layer referred to as 4BS , but reassessment of the sediments imply that it may have derived from 5BS (Beaumont et al. 1978; Cooke et al. 1945; Griin and Beaumont 2001; Grun and Stringer 1991). Nitrogen analysis is used to suggest that Border Cave 1 and 2 are of a similar age, yet their values differ from those of faunal samples that are supposedly from the same level (Beaumont et al. 1978).

Border Cave 1 (de Villiers 1973) consists of a fragmentary cranium including much of the frontal squama and supraorbital region, portions of the parietal and temporal bones and an occipital fragment. The only portion of the facial skeleton preserved is a part of the right zygomatic bone. The cranium appears to have been relatively long, broad and moderately high. The vault is quite robust , especially in the mastoid region and the occipital. There is little postorbital constriction and only moderate parietal bossing . The supraglabellar segment of the broad frontal is very steep, but behind this region the frontal recedes gently towards the bregma. The supraorbital region is moderately developed , but does not form a torus in that it is absent over the nasal root. It is difficult, however , to separate the superciliary arch from the supraorbital trigone. The interorbital region is quite broad and the nasion is depressed. There is a ridge on the malar of Border Cave 1 that may be a malar (zygomaxillary) tuberosity .

Border Cave 3 was recovered from what was described as a "shallow grave" cut down into the 4BS layer from, at the earliest, the final Middle Stone Age (Epi-Pietersburg) levels, (Beaumont 1980; Beaumont et al. 1978; Grun and Beaumont 2001; Griin and Stringer 1991). A perforated Conus shell was found in association with Border Cave 3. This shell and one from the lRGBS.UP layer have similar amino acid analyses that has been interpreted as indicating that Border Cave 3 was buried at the time of the deposition of the IRGBS layer (Grun and Beaumont 2001). The Conus shell has also provided a radiocarbon date of 33,570+/-120 years BP (AA14033) that is thought to be a minimum age for Border Cave 3 (Griin and Beaumont 2001). Border Cave 5 is reported as having been eroding out of a Middle Stone Age 2 (Middle Epi-Pietersburg) level 0.25m below the surface of the 3WA layer abutting a shallow depression (Beaumont 1980; Beaumont et al. 1978; Griin and Beaumont 2001; Griin and Stringer 1991). The 1.8m wide semi-circular depression was subsequently excavated but did not produce any further human remains (Griin and Beaumont 2001). Beaumont (Grun and Beaumont 2001) has argued that sedimentology and artefact analysis indicates that Border Cave was recovered from undisturbed layers and so could not have been an Iron Age burial.

Border Cave 2 (de Villiers 1973) comprises most of the corpus and the anterior borders of the rami of an adult mandible without teeth . The corpus is of moderate height and robusticity , while the anterior surface of the symphyseal region presents a well-developed mental eminence. Border Cave 2 appears to be too narrow and low to have belonged to the Border Cave 1 cranium . Border Cave 3 (de Villiers 1973) is the fragmentary skeleton of an infant aged between four to six months. The disarticulated skeleton, which was recovered from a shallow

44

Nitrogen analyses are used to support the dating of Border Cave 3 and 5 as younger than Border Cave 1 and 2 (Beaumont et al. 1978).

conjecture. Nitrogen analyses have been used to demonstrate that the hominids are not intrusive into the levels they are associated with (Beaumont 1980; Beaumont et al. 1978), yet there are variations in the values for Border Cave 1, faunal material from the same levels, and for the hominids and fauna from supposedly the same levels. These anomalies are explained as being the result of the heating of the faunal samples by overlying and/or contiguous hearths, something that is said not to have affected the hominid samples, and because the fauna and hominid bones underwent nitrogen depletion at different rates (Beaumont et al 1978). This does not explain why two different laboratories provide markedly different nitrogen values for Border Cave 1 (Beaumont 1980; Oakley et al. 1977). The explanations for the differences are not convincing and may explain the use of the term "most likely interpretation" in relation to the results of the nitrogen analyses (Beaumont 1980; Beaumont et al. 1978).

McBrearty and Brooks (2000) cite a preferred age estimate of 90-115,000 years BP for the Border Cave hominids based on geomorphology, associated fauna, amino acid racemisation and extrapolation from overlying radiocarbon dates. There are problems with the dating of the Border Cave material. The association of the hominid material with specific levels remains open to debate as only Border Cave 3 was recovered during systematic archaeological excavation and it could be intrusive (Border Cave 5 was found eroding out of a section). The postcranial material was recovered from sediments that had slumped into the 1941-2 excavations and so the exact stratigraphic context is not known (Morris 1992).

It is also significant that the hominid remains are better preserved than the faunal material from the supposedly equivalent levels, which is highly fragmented as a result of substantial post-depositional leaching and profile compaction (Klein 1983). Beaumont (1980) attempts to explain this difference by suggesting that all of the hominid material may have been from Middle Stone Age burials. If this suggestion is correct, it would make Border Cave even more anomalous within the Middle Stone Age of sub-Saharan Africa.

The estimated ages for the Middle Stone Age levels at Border Cave and the inferred dates for the hominid remains were initially based on sedimentological analyses coupled with the radiocarbon framework for the upper levels at the site, which were used to correlate the stratigraphy with oxygen isotope stages 4, 5 and 6 (Butzer et al. 1978). This correlation was used to give the basal Middle Stone Age deposits an age of approximately 195,000 years BP, placing the hominids between 90,000 and 115,000 years BP (Beaumont 1980; Butzer et al. 1978). As no direct correlation with sequential changes in the isotopic composition of sea water can be shown at this inland site, these dating estimates involve assumptions that relegate them to reasoned guesses (Parkington 1990).

Parkington ( 1990) has questioned many of the assumptions inherent in the dating of both the hominids from, and strata at, Border Cave, and highlights the disagreement in the dating of the lower levels between Beaumont and Butzer (Beaumont 1980; Beaumont et al. 1978; Butzer et al. 1978). Griin (Grun and Stringer 1991; Griin et al. 1990) has established an electron spin resonance (ESR) chronology for the stratigraphy and archaeological sequence at Border Cave. It should be noted that the large mammalian teeth used for the ESR dating did not contain any significant amounts of uranium, which is unusual considering teeth and bones show post-depositional uranium uptake. The ESR results showed internal stratigraphic consistency (excluding two anomalies). The brown sand layer 4BS, from which Border Cave 1 and 2 may have been derived and into which Border Cave 3 was apparently buried, has age ranges from 62-85,000 years BP, whereas the white ash layer 3WA from which Border Cave 5 was purportedly extracted, has ages ranging from 50-65,000 years BP (Griln and Stringer 1991:Figure 26; Griln et al. 1990:Fig. 1). It is suggested that if the inferred stratigraphic positions of the hominids are accepted, Border Cave 5 would date to 50-65,000 years BP, Border Cave 3 to 70-80,000 years BP and Border Cave 1 and 2 to greater than 90,000 years BP (Griin and Stringer 1991).

Even if the deposits are old, it is not necessary for the hominid material to be of a similar age. The results of nitrogen analyses are used to show that Border Cave 3 was not intrusive, yet the value for the hominid is reasonably different from that of faunal bone from the same level (Beaumont 1980; Beaumont et al. 1978). Cook et al. ( 1945) and Beaumont ( 1980) have concluded that the grave for Border Cave 3 would have been very shallow, with a depth of only 24cm and the grave lip marginally above the first of the bones. Sediments identified as overlying Border Cave 3 have been dated to 105,000 years BP from amino acid racemisation calibrated with radiocarbon dates higher in the sequence (McBrearty and Brooks 2000; Miller et al. 1999), but there results need further confirmation. If the attribution of Border Cave 3 to Middle Stone Age levels is accepted, it would be the only example of a burial from this period in sub-Saharan Africa (Volman 1984). The associated perforated Conus shell pendent would also be the only such example dated to the Middle Stone Age period in subSaharan Africa (Volman 1984). Finally, the use of red ochre in burial practices only becomes a common practice during the Late Stone Age in sub-Saharan Africa (Deacon 1984). These points suggest that the grave for Border Cave 3 may have been dug from a higher, possibly early Late Stone Age, level. It has also been suggested that the diagenetic condition of the bones of Border Cave 3 are not consistent with a very early date (Sillen and Morris 1996).

Griin (Griln and Beaumont 2001:Figure 5, Table 2) has revised this ESR chronology with the ESR age estimations up to 30% older. The following revised mean ESR ages have been calculated for the following layers: Layer 3WA66,000+/-2,000 years BP (BC5), lRGBS-76,000+/-4,000 years BP (BC3), 4BS-82,000+/-2,000 years BP (BCl), 5BS166,000+/-6,000 and 147,000+/-6,000 years BP (BCl) (Griln and Beaumont 2001 ). These revised ages mean that Border Cave 5 dates to around 66,000 years BP, Border Cave 3 to around 76,000 years BP and Border Cave 1 and 2 to around

The contextual problems that surround Border Cave 1 and 2 and possibly Border Cave 5 make their dating pure

45

either 82,000 years BP or 170,000 years BP (Griin and Beaumont 2001).

Van Vark, Bilsborough and Dijkema (1989) used various multivariate statistics, including Mahalanobis' distance , to compare Border Cave 1 with recent population samples, a late Upper Palaeolithic and Epipalaeolithic sample, an early Upper Palaeolithic sample , a Neanderthal sample and a Homo erectus sample. They found that, overall, Border Cave 1 was more different from the recent population samples than were the Upper Palaeolithic samples (van Vark et al.1989). They suggested that Border Cave 1 was not as modem as many authors contend. Corruccini ( 1992) reached a similar conclusion.

The Howieson's Poort layers at Border Cave were initially given ages of between 85-95,000 years BP (Beaumont et al. 1978), but have also provided much younger ESR ages of between 45-75,000 years BP (Griin et al. 1990). Revised mean ESR ages now place the Howieson's Poort layers at between 60-79,000 years BP (Griin and Beaumont 2001). The Howieson's Poort levels at the sites of Border Cave, Boomplass and Apollo 11 have also been bracketed by "limiting dates" of 56,000 and 80,000 years BP centred around 66,000+/-5,000 years BP, based on the extent of isoleucine epimerisation in ostrich eggshells (Miller et al. 1999). These age ranges are consistent with ESR results from Klasies River Mouth (Griin and Stringer 1991), but differ to results •obtained from other methods used at Klasies River Mouth .

Morphologically , Border Cave 1 displays a mosaic of archaic and more modem features. The vault is relatively robust , especially in the masto id and occipital regions . The supraorbital region is moderately developed, but does not form a torus and it is difficult to clearly separate, the arch from the trigone. This latter feature is reminiscent of the earlier more robust hominids such as Kabwe 1, Saldanha I and Bodo 1. The broad frontal and slight degree of postorbital constriction of Border Cave 1 is similar to that of Florisbad, while the parietal bossing and the steep supraglabellar region recall Laetoli HI 8. In overall morphology , however , Border Cave 1 is generally more modem than these other hominids .

Recent direct dating rejects the postulated 90,000 to 115,000 years BP date for the Border Cave hominids. The "splitting factor" of hydroxyapatite samples from Border Cave 3 and Border Cave 5 indicate ages of less than 20,000 years BP (quoted in Pearson and Grine 1996). Results for the postcranial material was consistent with MSA fauna from the site, suggesting an age in excess of 40,000 years BP (quoted in Pearson and Grine 1996). Low levels of nitrogen content were recorded for the postcranial material, which is also consistent with MSA fauna from the site (quoted in Pearson and Grine 1996). These dates need to be confirmed by other methods and direct age estimates obtained for Border Cave 1 and 2 before the dating of the Border Cave material can be resolved.

The Border Cave 1 cranium is generally consistent with the morphology of the other hominid remains from Border Cave, which are essentially modem in their configurations (de Villiers 1973, 1976). On the analysis of mandibular nonmetric traits (Table 1:4, Fig. 5:16) Border Cave 2 grouped with European Upper Palaeolithic mandibles, while Border Cave 5 grouped with Klasies River Mouth KRM 41815 and Abri Pataud 1. The mandibular material is similar to some of the other mandibles from Klasies River Mouth, which are also essentially modem and have well-developed mental eminence.

The state of preservation of the hominid remains and the direct dating evidence suggests that the cranial and mandibular material, at least, are from later Middle or Late Stone age contexts.

The Border Cave ulna, is similar in size to that from Klasies River Mouth and the individual measurements can fit within modem samples. However , on a canonical variates analysis it was, along with the Klasies River Mouth ulna, at the margin of the 95% density ellipse of the Epipalaeolithic sample from Jebel Sahaba and modem samples (Pearson and Grine 1996). This placement of both the Border Cave and Klasies River Mouth ulnae reflects an archaic pattern combining a relatively high olecranon process with a low coronoid process (Pearson and Grine 1996). The thick cortical bone of the Border Cave humerus could also be an archaic feature.

To include Border Cave 1 in analyses of a reasonable number of measurements it would have been necessary to include many dubious estimated values (cf. de Villiers 1973 who provides measurements for Border Cave 1, over half of which are questionable) so it was not analysed during the current investigation. In earlier analyses Border Cave 1 was found to group with Steinheim, Skhul 5 and Jebel Qafzeh 6 (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). In other analyses Omo 1 and Florisbad also grouped with these hominids and so displayed similar relationships to Border Cave 1 (Habgood 1982). In a principal components analysis of the frontal squama carried out by Brauer (1984a) , Border Cave I was well separated from both archaic and anatomically modem hominids , and was on the opposite side of the graph to Saldanha 1 and Kabwe 1.

Stringer et al. ( 1979) placed the Border Cave hominids , along with the Klasies River Mouth material, in their Homo sapiens Grade 3, while Brauer (1984a) placed them in his Homo sapiens Grade 3 ([early] anatomically modern Homo sapiens) with Omo 1 and the material from Klasies River . McBrearty and Brooks (2000) placed the Border Cave material, along with Omo I and the Klasies River Mouth material into their Group 3.

Using Mahalanobis' Generalised Distance statistic , de Villiers ( 1973) found Border Cave 1 to be closest to the late Pleistocene Tuinplaas (Springbok Flats) cranium , but the other material in the analysis consisted of only recent African populations , and so all that the analysis demonstrates is that Border Cave and Tuinplaas differ from modem populations , and not their overall similarity .

Because of the robust but essentially modem morphological pattern of the Border Cave hominids , and their suggested early date, the site is very important for discussions of hominid evolution. As outlined earlier, the dating of the site, and so the position of the hominid material in the evolutionary sequence, are not secure and could be decidedly

46

1989:Fig. 6). There is no stratigraphic connection between Cave 1 and Shelter lA and Shelter IB, but the stratigraphy has been correlated by: ESR measurements on shell from layer 10 in Shelter 1B and layer 37 in Cave 1 group together that is suggestive of a similar age (Goede and Hitchman 1987); and The occurrence of ostrich eggshell that does not occur in the LBS member, but is found in layer IO in Shelter 1B and SCBl-2 in Shelter IA.

younger than is presently suggested. These dating problems mea~ th_atit is difficult to make firm conclusions regarding the sigmficance of the Border Cave skeletal material.

Klasies River Mouth Hominid remains have been recovered from Klasies River Mouth (KRM), a complex of caves (Caves 1 and 2 and Shelters lA, 1B and C) on the Tsitsikama coast, South Africa (Fig. 2:1; Deacon and Geleijnse 1988; Deacon et al. 1986; Singer and Smith 1969; Singer and Wymer 1982). These caves have a long Middle and Late Stone Age archaeological sequence (Singer and Wymer 1982; Wurz 2002).

Studies of the sedimentary sequences (Butzer 1978; Deacon and Geleijnse 1988), oxygen isotope analysis of shell samples (Deacon 1995; Deacon and Geleijnse 1988; Shackleton 1982) and Quaternary sea levels (Hendey and Volman 1986) place the earliest Middle Stone Age (MSA) ?ccupation of KRM within the last interglacial (oxygen isotope substage 5e), at approximately 120,000 years BP. This placement is consistent with the mammalian fauna (Klein 1974, 1976, 1983), micromammals (Avery 1987) and molluscan material (Voigt 1982) from the earliest levels at ~e site. MSA 2 levels have been placed within oxygen isotope substage 5c or 5a (Shackleton 1982). Further studies have confirmed the correlation of the LBS member with oxygen isotope substage 5e and have suggested that the SAS ~ember may correlate with the transition from oxygen isotope substage 5a to stage 4 (Griin et al. 1990; Rightmire and Deacon 1991).

One of the major goals of the original excavations at KRM was to recover a sample of Late Pleistocene hominid remains (Singer and Wymer 1982) and less attention was given to contextual details. A research program being conducted at KRM by H.J. Deacon (1989; 1995; Deacon and Geleijnse 1988; Deacon et al. 1986) has attempted to improve this situation so as to place new and old finds more securely within the stratigraphy at the site. The hominid skeletal material from KRM is generally very fragmentary and much of it provides only limited morphological information. The incomplete and fragmentary condition of the material suggests carnivore activity, although some of the bones have been burnt which opens up the possibility of interpersonal violence and/or cannibalism (Singer and Wymer 1982; White 1987). Only the most informative hominid specimens will be discussed. Descriptions of the other hominid remains can be found in Singer and Wymer (1982), Rightmire and Deacon (1991, 2001), Brauer et al. (1992), Grine et al. (1998), Churchill et al. (1996) and Pearson and Grine (1997).

Amino acid racemisation values for bone from the MSA levels in Cave I have provided a sequence of dates ranging back to 110,000 years BP for the earliest levels with human occupation (Bada and Deems 1975), although there are uncertainties in calibration with these dates (H.J. Deacon pers. comm.; Rightmire and Deacon 1991). One phase of stalagmite formation in Cave 1 occurred after the deposition of Singer and Wymer's (1982) MSA 1 levels and prior to the deposition of their MSA 2 levels (broadly LBS and SAS members of Deacon). This interval has been dated by the uranium disequilibrium method to between 110,000 and 98,000 years BP (Deacon 1989; Deacon et al. 1986). Uranium-series results on speleotherms suggest that the LBS member has a minimum age of 110,000 years BP (Griin et al. 1990; Rightmire and Deacon 1991).

Studies continue at KRM so as to obtain a more refined dating of the stratigraphy at, and finds from, the site (Butzer 1978; Deacon 1985a, 1989, 1995; Deacon and Geleijnse 1988; Deacon and Shuurman 1992; Deacon et al. 1986; Deacon et al. 1988; Griin and Stringer 1991; Griin et al. 1990; Shackleton 1982; Singer and Wymer 1982). As Deacon states These are truly complex deposits and the complexity has been increased by slumping and slope movement. (l 985a:59) Deacon (Deacon and Geleijnse 1988; Deacon et al. 1986) has divided the · KRM stratigraphy into a number of broad members that have been related to the layers identified by Singer and Wymer (1982). In broad terms, the oldest member consists of light brown sands (LBS member - layer 38 in Cave I & layer 15 in Shelter 1B - MSA I), followed by a thick sands-ash-shell member (SAS member - layers 37, 17-14 in Cave 1 and 39-23 in Shelter lA & above layer 14 in Shelter 1B - MSA 1 [lower part] & MSA 2), a rockfall member (RF - layer 22 in Shelter lA - MSA 2) and an Upper member (UM - layers 21-1 in Shelter 1A Howieson' s Poort, MSA 3)

The results from the various methods detailed above indicate that the initial occupation of Cave I at K.RM can be placed into the last interglacial. Shelter IA contains a similar, although much more complete stratigraphical record to that from Cave 1 (see Deacon [1989] for a detailed description and discussion of the stratigraphy). The basal sand units have been equated with a high sea level during oxygen isotope stage 5e (Deacon 1989; Shackleton 1982), whereas the end of the MSA occupation in Shelter IA is estimated to date to 60,000 years BP (Deacon 1989; Deacon et al. 1986). MSA 2 levels in Shelter lA are thought, on the basis of uranium disequilibrium results, to be younger than 100,000 years BP (Deacon I 989). Hendey and Volman (1986:194) have suggested that the entire MSA occupation of K.RM was "largely, or entirely, confined to the last interglaciation", although most other scholars would extend the occupation down into the last glacial (Binford I 984, 1986a; Butzer 1978; Deacon I 985a,

The LBS and overlying SAS members at the site are separated by a discontinuity of unknown duration. In Cave 1 the discontinuity is marked by layer 37/SAS RBS, whereas in Shelter lA it is labelled SCB 1-2 (Deacon and Geleijnse

47

1989; Deacon and Geleijnse 1989; Deacon et al. 1986; Shackleton 1982; Singer and Wymer 1982; Thackeray 1986, 1988). Much debate has surrounded the dating of Shelter 1B in general, and the modem appearing mandible KRM 41815 in particular. Shelter lB has been equated with the earliest levels of human occupation in Cave 1 even though there is no stratigraphical connection between them. Singer and Wymer ( 1982) based this correlation on the occurrence in both deposits of flake blades with abrasion of the striking platform, and the presence of a pebble beach at the base of the stratigraphic sequence in Shelter 1B and Cave 1. However , blades with abraded striking platforms are also found in MSA 2 levels in Shelter IA (Singer and Wymer 1982). There are also problems with the correlation of the two pebble beaches. Deacon (1989) argues that the oldest MSA levels in Cave 1 have no direct relationship to any beach deposit. Butzer ( 1978) has suggested that the pebble beach in Shelter 1B could have been laid down at the same time as the last of the MSA 2 levels in Cave 1 (Layer 14), which he has identified as a storm beach . This suggestion has not been accepted by all, with Singer and Wymer (1982) concluding that Layer 14 was formed by material rolling down from a sloping mound of occupational debris outside the cave mouth, and Hendey and Volman ( 1986) suggesting that the material was transported from outside the cave by water from torrential rains. Singer and Wymer (1986) have reaffirmed their earlier dating of Shelter lB.

supports my contention that Cave IB is most likely referable to the era represented by Howiesan's Poort and later deposits. (1986b :515) Deacon (1985a, 1989) argued that Shelter 1B has deposits that are as old as those in Cave 1, but that the mandible KRM 41815 was recovered from just above a discontinuity that separated the oldest and more recent levels in the shelter. This discontinuity could explain the difference in the oxygen isotope value for the single shell from Shelter 1B and those from the oldest MSA level in Cave 1 (Binford 1984, 1986a; Deacon 1989; Shackleton 1982). Relative dating of Turbo shells by electron spin resonance (ESR) analysis found that a sample from Shelter lB , which came from just below the mandible and apparently above the disconformity (Deacon 1989), was closest to MSA I samples , and not MSA 2, MSA 3 or Howieson's Poort samples (Goede and Hitchman 1987). The Shelter 1B sample, however, was distinguished from the samples from the oldest MSA levels in Cave 1, which suggests it is younger (Deacon 1989; Goede and Hitchman 1987). Deacon ( 1989) contends that the correlation of the Shelter 1B deposits above the disconformity to oxygen isotope stage 5b rather than 5e is suggested by these finds, and that an age closer to 90,000 years BP, rather than 100,000 years BP or older, is more acceptable. In regard to mandible KRM 41815 from Shelter 1B Deacon ( 1989) suggested that any debate rests on its morphology and not its date.

Binford (1984, 1986a, 1986b) has questioned Singer and Wymer's (1982) correlation of Shelter 1B with the earliest human occupation of Cave 1. He suggested that the contents of Shelter lB were more comparable to later levels at KRM, and not with the MSA 1 levels. Binford based this suggestion on core forms, the size of flakes, ungulate and molluscan species grouped by ecotype, faunal species abundance shifts, and on the oxygen isotope composition of a shell from Shelter IB. All of these form a regular pattern or sequence throughout the stratigraphic succession at KRM. Binford (1984, 1986a) felt that in these aspects Shelter 1B fitted more parsimoniously between Singer and Wymer ' s (1982) Howieson's Poort and MSA 3 levels, rather than with the MSA 1 levels of Cave 1. He suggested that ...caution should be exercised when considering Shelter 1B as contemporaneous with the early levels of Cave 1. (1984:41)

Grun (Grun and Stringer 1991; Grun et al. 1990) has also obtained ESR ages for the MSA levels at KRM. The ESR results reflected the stratigraphic positions of the samples used. The mammalian tooth fragments used contained significant amounts of uranium resulting in considerable differences in the ages from early uptake (EU) and linear uptake (LU) results. LU ages of 94,000 +/-10 and 88,000+/8,000 years BP for a sample from the lower SAS member is in reasonable agreement with uranium series dates detailed above, which prompted Griin to suggest that the LU results are more likely to be correct and that the EU results were gross underestimates (Grun and Stringer 1991; Griin et al. 1990). If the LU results are more acceptable, the earliest hominids from the LBS member at KRM would be older than 90,000 years BP , whereas the other material from the SAS member would date to between 60-85,000 years BP (Grun and Stringer 1991:Figure 24; Grun et al. 1990: Fig. 3). However , if the EU dates were correct , the hominid material would date to less than 80,000 year BP (Griin and Stringer 1991:Figure 24; Grun et al. 1990: Fig. 3).

Thackeray (1986) has challenged Binford's proposal in relation to the fauna from Shelter lB (see also Deacon 1985a). Thackeray (1986) carried out a factor analysis of the relative abundance of ungulate species and found that the Shelter lB fauna! assemblage was most similar to that from MSA 2 levels in Cave 1. Thackeray (1986:511) does, however, caution that "similarity in faunal indices does not necessarily imply contemporaneity ". In reply, Binford ( 1986b) performed his own multivariate analyses (principal components and cluster analyses on normalised data), in which the Shelter 1B sample was most similar to MSA 4, MSA 3 and Howieson's Poort samples. Binford (1986b:515) felt that these results were "in the same ballpark " as his earlier assessments, and stated ...Thackeray's analysis is pro cedurally flawed and that a more informed analysis [his own?]

McBrearty and Brooks (2000) cite preferred age estimates of 118,000 years BP for the LBS member and 94-105,000 years BP for the SAS member . There remain problems with the dating of the KRM deposits. For example , on sedimentological grounds Butzer (1978) placed the Howieson's Poort levels at around 95,000 years BP, Shackleton (1982) , using oxygen isotope analysis, favoured a date between 50,000 and 30,000 years BP although he did not exclude an earlier date, whereas Deacon (1985a) can see no good evidence for dating them younger than 65,000 years BP. Griin (Grun and Stringer 1991; Griin et al. 1990) obtained ESR LU ages in the 40-60,000 years BP

48

range for the Howieson's Poort levels, and even younger with EU ages. Border Cave has provided initial ESR ages of between 75,000 and 45,000 years BP for Howieson's Poort levels (Grun et al. 1990), which have been revised and now place the Howieson's Poort layers at between 60-79,000 years BP (Grun and Beaumont 2001). Also, the Howieson's Poort levels at the sites of Border Cave, Boomplass and Apollo 11 has been bracketed by "limiting dates" of 56,000 and 80,000 years BP based on the extent of isoleucine epimerisation in ostrich eggshells (Miller et al. 1999). This demonstrates how difficult it is to determine the age of the Howieson's Poort levels at Klasies and comparable sites with estimates ranging from 30-95,000 years BP. Parkington ( 1990) has discussed in detail the problems that surround the dating of KRM.

SAM-AP 6269 [layer 16] is a right temporal fragment preserving a glenoid fossa, root of the zygomatic arch, a small section of the tympanic plate and part of the anterolateral surface of the mastoid process (Grine et al. 1998). The form and size of the temporal fragment suggest that the individual would have been comparatively small. The glenoid fossa is described as reasonably small, but moderately deep with a distinct articular eminence and a small postglenoid process. The posteromedial wall of the glenoid fossa is composed entirely of the squamous temporal without contribution from the tympanic (Grine et al. 1998). KRM 27889 [layer 15] is the proximal end of a lightly constructed left radius preserving the head, neck and tuberosity. The cortical bone of KRM 27889 is moderately thick and the neck is relatively stout (Pearson and Grine 1997). Like other Klasies material, this fragment is suggestive of a small individual. KRM 16424 [layer 14+ or a disturbance in layer 15] consists of part of the right ramus and corpus, containing three small molar teeth, from a very small and gracile mandible. Whilst there is not a retromolar space, the leading edge of the ascending ramus is slightly posterior of the 3rd molar. Discolouration and bone flaking indicate that KRM 16424 was burnt. Singer and Wymer (1982) suggested that if the 3rd molar was not present, this specimen would have been identified as immature. KRM 16424 may have belonged to a small adult female. Sam-AP 6268 [layer 14] is an atlas vertebra fragment including parts of the anterior arch and lateral mass. KRM 26076 [layer 14] is part of a small lightly built left clavicle. KRM 16651 [layer 14] is a relatively large incomplete left zygomatic with a big zygomaticofacial foramen and a columnar-like lateral orbital pillar. Singer and Wymer (1982) described KRM 16651 as being flat in facial aspect. KRM 14695 [layer 14] consists of the anterior portion of a mandible without teeth. There is some bone erosion on the vertical symphysial surface, but the mental eminence, whilst present, would not have been markedly developed. KRM 13400 [layer 14] is a mandibular fragment, including much of the right corpus with teeth, and the symphyseal region. The fragment is quite robust, with a prominent mandibular torus on the basal portion of the corpus. There are mental foramen positioned under the M 1 and the PM4. The symphysis region is almost vertical with a slight mental protuberance, but it does not have the central keel or thickened inferior margin that Schwartz and Tattersall (2000) argue is typical of modem humans.

Whilst Deacon ( 1985a, 1989; Deacon and Geleijnse 1989; Deacon et al. 1986) would argue for older ages for the KRM levels and the associated hominid material, taking into consideration all of the available evidence a reasonable estimate would place the LBS member at 90-100,000 years BP and the SAS member at between 60-85,000 years BP. Deacon et al. stated The Klasies River Mouth sites when adequately dated will serve as the key reference for the cultural stratigraphy of the Middle Stone Age in the southern Cape and further afield. (1986:36) The site will also serve as a key to explaining the timing of the appearance of anatomically modem humans in subSaharan Africa.

Cave 1 No diagnostic human remains come from the LBS member within Cave 1, whereas the SAS member has produced the bulk of the hominid material from the site. The following are the most diagnostic material recovered. Detailed descriptions of the hominid finds from Cave 1 are provided by Grine et al. ( 1998), Pearson and Grine ( 1997), Rightmire and Deacon (1991, 2001) and Singer and Wymer (1982). KRM 27070 [layer 37] is a thin parietal fragment. KRM 2177 6 [layer 17] is a relatively large left mandibular corpus without teeth. A large mental foramen is positioned below the PM4-Ml. A chin is represented by a moderately developed trigone, but it may appear more prominent due to slight alveolar prognathism. The surface of this specimen is blackened suggesting that it may have been burnt. KRM 16425 [layer 16] is a frontal fragment consisting of the glabellar region, part of the right supraorbital and the frontonasal suture and broken nasal bones. The glabella is moderately inflated, but the superciliary segment is only slightly developed. The nasal root is flat and relatively broad, while the superior orbital margin is slightly rounded in shape. KRM 16425 is essentially modem in form, with a gracile supraorbital region, although the interorbital area is quite broad. The degree of frontal sinus development suggests that this fragment is from an adult individual, but a younger age cannot be excluded, with Frayer et al. (1993, 1994) suggesting that KRM 16425 may be from a juvenile.

Dental remains from the lower SAS member (sub-member SAS U, unit SMB) in Cave 1 confirm that some of the individuals from the site had quite small molars (Rightmire and Deacon 2001 ).

Shelter lA Limited hominid remains have been recovered from Shelter IA (Brauer et al. 1992; Churchill et al. 1996; Deacon 1989; Rightmire and Deacon 1991; Singer and Wymer 1982). The LBS member has produced two fragmentary maxillae fragments found in the same shell-rich midden heap within a

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developed mental eminence that may be accentuated by the resorption of the alveolar margin. Schwartz and Tattersll (2000) have described KRM 41815 as having a mental protuberance that tapers superiorly into a central keel and expands inferiorly into bilateral mental tuberosities, a morphology closer to that of modem humans than the other mandibular remains from the site. In overall morphological configuration this mandible is essentially modem in form. KRM 41820 [layer 1O] is a moderately robust mandibular condyle that was found near KRM 41815 and may belong to its right ramus (Singer and Wymer 1982).

metre of each other (Brauer et al. 1992; Rightmire and Deacon 1991). AA43/SAS4SHB is a burnt left edentulous maxillary fragment broken along the intermaxillary suture retaining part of the left alveoli and a portion of the hard palate. This specimen, which may be from a young individual, is quite robust and the palate is quite deep . the morphology of AA43 has been described as being similar to that of Kabwe 1, although smaller in size, (Deacon 1989), and in overall size and proportions to Laetoli H 18 (Rightmire and Deacon 1991). Brauer et al. (1992:421-2) concluded that measurements of AA43 do not provide evidence of any "pronounced robusticity that could indicate an archaic morphology". Z44/SAS4SHC is a small fragment of a left maxilla with a small 1st molar in place. 244 has been described as not only gracile when compared to AA43 , but also when compared to modem human standards (Rightmire and Deacon 1991).

The Klasies River Mouth hominid material was too fragmentary to include in any metrical analyses. On the analysis of mandibular non-metric traits, the mandible from Shelter lB, KRM 41815, grouped with Border Cave 5 and Abri Pataud 1 (Table 1:4, Fig. 5: 16). Pearson and Grine ( 1997) undertook canonical variates analysis of six measurements from the left radius KRM 27889 comparing it to a recent human , "early modem" [Omo 1, Skhul 2, 4, 5 & 7 & Jebel Qafzeh 9] and European and western Asian Neanderthal samples. On a plot of the first two canonical axes, they found that KRM 27889 was placed outside the 95% confidence ellipse of the recent human sample, near the outer edge of the Neanderthal ellipse, but well within the "early modem" ellipse (Pearson and Grine 1997:Figure 2). It is worth noting that a radius from the Cave of Hearths included within the analysis fell within the 95% confidence ellipse of the recent human sample .

The two maxillae fragments have been interpreted as indicating that there was a significant degree of sexual dimorphism evident within the Klasies population during the time of the LBS member within Shelter IA Hl8 (Rightmire and Deacon 1991). From the SAS member: KRM 41658 (layer 36) is a relatively thick and robust parietal fragment. Singer and Wymer (1982) noted that the form of this specimen suggested a low and narrow vault, whereas Rightmire and Deacon ( 1991) did not think that the vault had to be especially low. KRM ulna consists of the proximal half of a gracile right ulna described as displaying a relatively low coronoid height and a reduced lateral flare at midshaft. Rightmire and Deacon ( 1991) concluded that this bone was similar in dimensions to modem humans, but this conclusion has been questioned by Churchill et al. ( 1996) who found the KRM ulna to be archaic in overall morphology and not fully modem

Grine et al. ( 1998) undertook canonical variates analysis of five measurements from the temporal SAM-AP 6269 comparing it to recent Khoisan and Zulu samples and four Pleistocene samples (Neanderthals, European Middle Pleistocene [Steinheim, Petralona & Arago 21 ], African Middle Pleistocene [Sale, Ndutu & Kabwe 1] and Pleistocene "early modern" [LH 18, JI 1, JI2, Singa, Omo 2 & Skhiil 4 & 5]. On a plot of the first two canonical axes, they found that SAM-AP 6269 fell within the 95% confidence ellipses of the Khoisan and Zulu samples, but it was near the outer edge (Grine et al. 1998:Figure 4). Whilst SAM-AP 6269 was placed nearer the modern samples on canonical axis 1, it was placed with the Pleistocene "early modern" African Middle Pleistocene hominids on canonical axis 2. The relatively small size of this specimen may be contributing to its placement within the modem African ellipses. Grine et al. ( 1998: 105) concluded that whilst the evidence is limited, it suggests that SAM-AP 6269 displays an "overall, but not unambiguous, pattern of modern morphology."

From the upper part of the MSA sequence of Shelter IA a number of isolated teeth were obtained. They are small and must have come from relatively gracile individuals (Deacon 1989).

Shelter lB Singer and Wymer (1982) equate the archaeological remains from this shelter with the lowest levels in Cave 1, which they classify as MSA 1 (see also Volman [1984] who classified these levels as Middle Stone Age 2), whereas Deacon (1989; Deacon and Geleijnse 1988; Deacon and Shuurman 1992) equates layer 15 to his LBS member and the deposits above layer 14 to his SAS member. KRM 41815 [layer 10] is a relatively complete mandible with damaged rami and missing anterior and some posterior teeth. The remaining posterior teeth are heavily worn, but would have had relatively small crowns. KRM 41815 appears to have had agenesis of the left third molar , which x-rays reveal is not unerupted. The right third molar is lost, but the socket is evident. The corpus is relatively short and quite robust. The tooth sockets indicate resorption of the alveolar margin. The ramus is broad and the coronoid process is large. The symphyseal region displays a well-

Churchill et al. ( 1996) found the KRM ulna to be intermediate between archaic and modem humans on a bivariate plot of olecranon height/coronoid height (Churchill et al. 1996:Figure 2) . It is worth noting that on the bivariate plot of olecranon height/coronoid height, ulnae from Skhul and Jebel Qafzeh were positioned nearer a modem human regression line than the KRM ulna (Churchill et al. 1996:Figure 2). Churchill et al. (1996) also undertook a canonical variates analysis of the KRM ulna from the SAS member of Shelter IA. On a plot of the 1st and 2nd canonical axes, the KRM ulna fell within the 95% density ellipse of an archaic sample (European and western Asian Neanderthals and Baringo 66 [Homo cf. erect us) and outside the 65%

50

density ellipse of a modem sample (Churchill et al. 1996:Figure 3). Churchill et al. (1996) conclude that, contra Rightmire and Deacon ( 1991), the KRM ulna was archaic in overall morphology and not fully modem. It is worth noting that on the plot from the canonical variates analysis, ulnae from Skhiil and Jebel Qafzeh are well separated from the KRM ulna, fall within the 65% density ellipse of the modem sample and are intermediate between the modem and archaic samples (Churchill et al. 1996:Figure 3). Similarities with the fragmentary ulna from Border Cave have also been identified (Pearson and Grine 1996). They both present an archaic pattern combining a relatively high olecranon process with a low coronoid process (Pearson and Grine 1996).

Limited metrical comparisons found the AA43 maxilla to fall within or close to the ranges for Holocene and sub-recent samples and well below the values for Kabwe I and, to a lesser extent, Laetoli H18 and Jebel lrhoud 1 (Brauer et al. 1992). Jebel Qafzeh 9 was closer to AA43 on some of the measurements. These results were viewed by Brauer et al. ( 1992) as confirming that AA43 was anatomically modem in overall morphology. Whilst Brauer et al. (1992:421) indicated that the growth of the anterior part of the maxilla would have been practically complete, even if the specimen had only been 14 years old, the possibility that the measurements were underestimating the adult dimensions of AA43 remains.

Wolpoff (1989) suggested that the superciliary arch on the frontal fragment, KRM 16425, was "not unlike" that of Florisbad, although Rightmire (1978a) and Singer and Wymer ( 1982) did not see similarities to Florisbad. Singer and Wymer (1982) also saw little similarity between KRM 16425 and Border Cave I. Wolpoff (Wolpoff and Caspari 1990) has also argued that the interorbital breadth of KRM 16425 exceeds that of Florisbad and Holocene samples from the southern Cape.

The hominid material excavated by Singer and Wymer at KRM indicate the simultaneous occurrence of gracile and robust forms. For example, the MSA 2 levels provided robust mandibles (KRM. 13400 and KRM 21776), as well as a gracile mandible (K.RM 16424), cranial fragments (KRM 16425 and KRM 16651) and gracile postcranial remains (KRM 26076). This morphological dichotomy did not stop Singer and Wymer (1982) from concluding that the KRM skeletal material demonstrated that Homo sapiens sapiens were in southern Africa prior to 100,000 years BP, and that they manufactured the early stages of the MSA.

It has been suggested that the size of the zygomatic KRM

16651 could only be matched within the African hominid sample on the Bodo face (Frayer et al. 1993, 1994; Wolpoff and Caspari 1990). Brauer (Brauer and Singer 1996; Stringer and Brauer 1994) has questioned the data on which this contention was based (the horizontal height measurement) and suggested that KRM 16651 falls within a modem range of variation. However, as Brauer (Brauer and Singer 1996) demonstrates, there are overall size and morphological similarities between KRM 16651 and the Florisbad and Laetoli H18 hominids. The columnar-like lateral orbital pillar is an archaic feature.

The more recent excavations at the site have confirmed this impression with the LBS member within Shelter IA producing two fragmentary maxillae fragments, one robust and one gracile, found in the same shell-rich midden heap within a metre of each other (Rightmire and Deacon 1991). Deacon (1989; Rightmire and Deacon 1991) has argued that the population occupying the site was typified by an extreme degree of sexual dimorphism. There is however, the possibility that much of the gracile hominid material could be from young individuals as no ages have been published for the various remains (see Frayer et al. 1993, 1994). Whilst this may account for some of the morphological variation evident at the site (eg. the small supraorbital development on KRM 16425 or small teeth), it cannot explain the very small mandible K.RM 16424 that has its 3rd molar in situ. As noted above, Singer and Wymer ( 1982) suggested that if the 3rd molar was not present, KRM 16424 would have been identified as immature. Mann (1995) has gone so far as suggesting that ... this mandible [KRM 16424] represents the first evidence of a little person (a midget) in the fossil record. (1995:41)

Lam et al. ( 1996) compared the KRM mandibular remains with a European and western Asia Neanderthal, Pleistocene sample early modem sample [Skhiil 4 & 5, Border Cave 2 & 5, Kish Hoek, Matjes River, Predmosti 3 and Cro-Magnon 1 & 2] and a recent human sample. On bivariate plots, the Klasies mandibles generally fell at the extreme or just outside the 95% density ellipses of the recent human sample. On the bivariate plot of chin indices, the Klasies mandibles are placed well outside the Pleistocene modem 95% density ellipse, but within or at the extreme of both the Neanderthal and the recent human 95% density ellipses (Lam et al. 1996:Fig. 6). These results indicate that the Klasies mandibles do not have well-development chins compared to the Pleistocene modem or recent human samples and even to some Neanderthals. Lam et al. (1996:554) contend that chin development on the Klasies mandibles is "much less pronounced than would be expected in extant populations."

Rightmire and Deacon have concluded that These studies confirm that the Klasies jaws, teeth and other body parts fall easily within the range of variation exhibited by modem humans. (1991: 132)

It is worth noting that on the bivariate plot of corpus height

and breadth at Ml, KRM 16424 falls outside the 95% density ellipses of the Neanderthal and recent human samples and at the lower extreme of the Pleistocene modem sample (Lam et al. 1996:Fig. 4), confirming that it is very small in overall size. Lam et al. ( 1996) contend that the small size of KRM 16424 is the most convincing evidence of the presence of modem humans at KRM.

However, there are differences between the morphological affinities of the more robust and the gracile individuals from KRM. For example: Singer and Wymer (1982) suggested that the robust mandibular remains were similar to the immature mandible from the Final Acheulean levels at the Cave of Hearths (Tobias 1971), which on faunal grounds appears to be close

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to, but slightly younger than, Kabwe 1 (Partridge 1982; Vrba 1982). The robust maxillary fragment AA43 is morphologically comparable to the maxillae of Kabwe 1 and 2, and similar in overall size and proportions to Laetoli H 18 and Jebel Irhoud 1 (Brauer et al. 1992; Deacon 1989 Rightmire and Deacon 1991; but also see Brauer et al. [ 1992] who found AA43 to fall within or close to the ranges for Holocene and sub-recent samples and well below the values for Kabwe 1). The size of the zygomatic KRM 16651 is comparable to either Bodo (Frayer et al. 1993, 1994) or the Florisbad and Laetoli H 18 hominids (Brauer and Singer 1996; Rightmire 1978a; Stringer and Brauer 1994) depending on the results of measurements (but see Brauer and Singer [1996] and Stringer and Brauer [ 1994] who suggest that KRM 16651 falls within a modem range of variation) . The KRM ulna is archaic in overall morphology (especially olecranon height/coronoid height) and not fully modem (Churchill et al. 1996).

still be the possibility of a chronological ordering within the KRM hominid sample . This scenario remains very speculative , and does not match with Deacon ' s views that are based on a detailed knowledge and understanding of the site. As noted above, the KRM hominid material has been described as being within the range of morphological variation of modem human samples. The limited postcranial material from the site suggests a more archaic morphological pattern . This is also evident from the morphological comparisons of much of the fragmentary material as detailed above . These comparisons suggest a morphological link with the hominids from Florisbad , Laetoli Hl8 , Border Cave (No. 5 and the postcranial material) and to a lesser extent Jebel Irhoud 1and Kabwe 1. Whils t the Klas ies material does have archaic elements , it has moved closer , morphologically , to modem humans than this earlier material. This could be either a chronological progression within the hominid material at the site or incorporated within an extremely sexually dimorphic and morpholo gically variable population . Stringer et al. (1979) placed the Klasies River Mouth material with the Border Cave hominids in their Homo sapiens Grade 3. Brauer (1984a) placed the Klasies River Mouth material , along with Omo 1 and the Border Cave hominids , in his Homo sapiens Grade 3 ([early] anatomically modem Homo sapiens ). Wolpoff and Caspari ( 1990:395) observed that "as a sample the Klasies material is not particularly modem" . Whilst the majority of the material is fragmentary and there is morphological variation within the sample, the KRM hominid material can best be categorised as representing archaic Homo sapiens. McBrearty and Brooks (2000) placed the Klasies River Mouth material , along with Omo 1 and the Border Cave material into their Group 3.

The gracile hominid remains, however , are essentially modem in their morphological configurations and affinities (Deacon 1989; Rightmire and Deacon 1991; Singer and Wymer 1982). Whilst the symphyseal region of most of the Klasies mandibles are generally vertical, the chin development is less pronounced than is generally indicated and much less than would be expected in modem populations (Lam et al. 1996). The KRM 41815 mandible displays general similarities with the mandibles from Border Cave, especially in the symphyseal region where there is a well-developed mental eminence . KRM 27889 left radius displays more similarities with archaic Homo sapiens material than to recent human samples (Pearson and Grine 1997). Right temporal fragment SAM-AP 6269 displays an overall pattern of modem morphology (Grine et al. 1998) KRM 16424 is a very small mandible when compared to both archaic and recent human samples , with Lam et al. ( 1996) suggesting that its small size is the most convincing evidence for the presence of modem humans at KRM, whereas Mann (1995:41) has declared that due to its small size "it is virtually impossible to asses its phylogenetic significance ". Z44 maxilla fragment has been described as not only gracile when compared to AA43, but also when compared to modem human samples (Rightmire and Deacon 1991 ). The size of the 1st molar is also extremely small (Lam et al. 1996).

Discussion The sub-Saharan African sequence of hominid fossils is generally regarded as documenting a local, in situ , evolution to some degree. The number of evolving lineages present in the region, and its/their relationship( s) to skeletal material from other areas has varied , depending on the number of crania available for consideration and the chronological scheme used. Hrdlicka ( 1930), considering only the material from Kabwe , thought that modem humans must have moved into Africa from Europe where he thought they had evolved . He suggested that Kabwe 1 represented ...a distinct crude variety of man, which strangely combines many ancient , even preNeanderthal conditions with others that are relatively modem. It could represent , conceivably , a very brutish individual development of the upper Neanderthal or the post -Neanderthal period . (1930 :124) However , he cautioned that Kabwe 1 was not a Neanderthal , and stated The Rhodesian skull is a tantal izing specimen to the student, who is wholly at a loss as to just where it belongs taxonomically or

Another possibility is that the apparent morphological dichotomy at KRM could indicate a chronological separation of the two morphological types , with the robust form being older than the more gracile type . This scenario would be supported if the mandible KRM 41815 from Shelter 1B is, as Binford (1984, 1986a, 1986b) suggested , considerably younger than the date assigned to it by Singer and Wymer (1982) or Deacon (1989) . However , the two maxillary fragments found within a metre of each other in a shell-rich midden within the LBS member of Shelter lA do not support such a scenario because they appear to be contemporaneous yet morpholo gically very differe nt. If Z44 was a young, small female yet to achieve adult dimensions there would

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chronologically. prehistory.

It is a comet of man's

Smith's conclusions for sub-Saharan Africa are similar to those of Wolpoff in that he contends From the standpoint of morphology, continuity is perceptible in many features ...and broadly speaking, each group is a suitable ancestral group for the next younger group. (1985:204) Smith et al. concluded Based on the mosaic of archaic and modem morphometric features exhibited by these specimens [Florisbad, Laetoli H 18, Omo Kibish ], they do appear to represent a logical transitional series between the more archaic hominids of the Kabwe group and early modem Africans. (1989:44)

(1930:130) Weidenreich (1947:201) concluded that "the morphological evidence suggest that man has evolved in different parts of the old world", and argued for a direct evolutionary relationship between Kabwe 1 and modem African groups via forms such as the Boskop calotte. Vallois argued that ...in Africa Rhodesian Man does indeed seem to represent, like the European Neanderthals, the end of a phylum which left no descendants, since Florisbad man, much closer morphologically to the Sapiens type, is chronologically anterior to Rhodesian man. (1958:495) He postulated that in sub-Saharan Africa, like Europe, there were two distinct lineages, one, consisting of Saldanha 1 and Kabwe 1, becoming extinct, and the other, as exemplified by Florisbad, evolving into modem man.

However, whilst authors like Wolpoff and Smith support local development of modem humans in Africa, they do not necessarily equate this with the actual origin of all modem Homo sapiens. Frayer et al. stated . .. there is clearly nothing in this material that would signal any kind of a close relationship with more modem humans in Europe and Asia ... [and] ... nothing in the African fossil record supports this continent as a unique motherland. (1993:36)

Coon ( 1962) saw two independently evolving lineages in Africa, one north and the other south of the Sahara. The southern lineage started with the Homo erectus fossil, Olduvai H9, and progressed via Saldanha 1, Kabwe 1, Cave of Hearths, and Border Cave 1. This line "became extinct by absorption into the ranks of the invaders" (Coon 1962:633). These invaders were Coon's northern line, who were forced south towards the end of the Pleistocene by an influx of people from western Asia into north Africa. Examples of this line outside of north Africa include the Singa calvaria, the hominid from Florisbad and the Boskop calotte. Coon did state that The African material, however, is less well documented [than that from Europe, Asia and Australia] and new conclusions may be reached as new evidence becomes available. (1962:659)

Wells (1969, 1972) identified local in situ evolution in subSaharan Africa, but instead of a single line he suggested at least two lines documenting a large morphological range. He stated that Kabwe 1 ...should properly be included in the subspecies Homo sapiens afer of Linnaeus. (1972:7) and regarded the Kabwe-Saldanha group ...as an early and probably abortive side branch of Homo sapiens a/er ... (1972:7) and ... far from being a primitive or ancestral H. sapiens a/er, appears on the contrary extremely and aberrantly specialized. (1969:172) According to Wells, the direct developmental line in subSaharan Africa was documented by the skeletal material from Omo, Border Cave, Klasies River Mouth, and possibly Florisbad. These fossils appear to fulfil Well's ( 1969: 172) proposition that "there awaits discovery in the later MidPleistocene of Africa a true ancestral Homo sapiens a/er".

As Coon thought, with additional skeletal material and a refined chronology, new explanations for the African material have been expounded. A number of scholars have followed Hrdlicka, Weidenreich and Coon in asserting local in situ evolution throughout the Old World (for example Smith 1985; Wolpoff 1980a), and so see an evolving lineage in sub-Saharan Africa. Wolpoff stated The South African evidence suggests a continuous line of hominid evolution, with early modem sapiens derived from the local archaic sapiens populations, as represented by Broken Hill, Saldanha, and the Cave of Hearths. ( 1980a:331) and that The late Pleistocene situation in East Africa is very similar to that in the southern part of the continent. .. (1980a:333)

After obtaining a series of dates for many of the southern African hominid fossils (most of which now appear much too young), Protsch (1975) divided the material into two lineages. One line (Homo sapiens rhodesiensis), made up of Saldanha 1, Kabwe 1 and the Cave of Hearths mandible, was said to be a side branch that developed some 50,000 years ago. The other line (Homo sapiens capensis ), which was also the ancestor of the former group and included Saldanha 2, Florisbad and Border Cave 1, supposedly developed into modem man (Homo sapiens a/er). Protsch concluded On balance, subSaharan Africa is the likely place of the origin of anatomically modem man ...

53

advent of the final MSA , that is, before 50,000 BC at the latest...but that they indeed manufactured the early stages of the MSA more than 100,000 years ago. (1982:149) while Beaumont stated that the Hominid remains from ...Border Cave ...and from the MSA I and II levels at Klasies River Mouth ...demonstrate the presence in southern Africa of anatomically modern Homo sapiens in Last Interglacial deposits dating to c. 90,000-130,000 B.P . (1980:32)

and that migration probably started into East Africa around 60,000 to 50,000 years ago and spread from there into other parts of the world . (1975a :316) After detailed analysis of the sub-Saharan African late Middle and earlier Upper Pleistocene skeletal sample (Rightmire 1984a, 1984b, 1989), Rightmire suggested It seems unlikely that southern Africa has witnessed any large scale replacement of its peoples during the last 100ky ( l 984a :321) and concluded ...it is increasingly likely that modem human s were established in southern Africa not long after the onset of the Upper Pleistocene ... ( l 984a:3 l 6) However , Rightmire was quite cautious about his conclusions and warned against blindly accepting the continuity of later Middle Pleistocen e population s with more recent groups. He advised that Information presently availabl e does not satisfactorily settle the question of whether Homo sapiens sapiens evolved gradually, in several parts of Africa , or whether anatomically modem people arose from a single source, somewhere in the sub-Saharan region or outside of Africa entirely. (1984a:316) He also stated that ...the Border Cave and Klasies materials do not to my mind prove that modem humans evolved first in sub-Saharan Africa. (1981 :21)

The relationships of the late Middle and earlier Upper Pleistocene sub-Saharan African crania evident from the results of the multivariate analyses and morphological comparisons are consistent with the explanations discussed above in that they demonstrat e morphological continuity within the sub-Saharan African region (Fig. 2:2). The crania from sub-Saharan Africa tended to cluster with each other, revealing a high degree of geographical identity and regional continuity . The skeletal material does not demonstrate , as Coon (1962) has suggested, a major influx of new and morphologically different peoples into the region, or, as Wells (1969, 1972) and Protsch (1975) proposed , the occurrence of two morphological types evolving alongside each other until one became extinct. However, as Brauer noted ...we must assume that a broad morphological spectrum existed in these regions [eastern and southern Africa] during the late Middle and early Upper Pleistocene. (1984a:389) This morphological spectrum has expanded since Brauer made this observation with revised chronological placements of some hominids such as Florisbad and the discovery of additional hominid material including that from Herto Bouri , Beret and Klasies River Mouth .

Brauer (1984a, 1984b, 1984c, 1989) has given a detailed explanation of the evolutionary sequence in sub-Saharan Africa. He proposed that archaic Homo sapiens , such as Bodo 1, Saldanha 1 and Kabwe 1, evolved into anatomically modem Homo sapi ens in southern and/or eastern Africa during the Middle Pleistocene. He argued that these anatomically modem humans had , by the early Upper Pleistocene , spread throughout much of Africa , as demonstrated by the hominid material from Klasies River Mouth, Border Cave, Omo and Singa . Anatomically modem forms are suggested to have then moved into north Africa, western Asia and Europe. Brauer concluded that The new reconstruction of the course of evolution in Africa and Europe ...make an African origin of Homo sapiens sapiens appear most likely. (1984c:1)

The late Middle and earlier Upper Pleistocene sub-Saharan African hominid material does display variation on a common morphological theme . It is possible to document a morphological progression from the earlier and more robust crania to the younger anatomically modem forms. As has been outlined, Brauer ( 1984a, 1989), using a "grade " model proposed by Stringer et al. (1979), divided the sub-Saharan African hominid sample into three grades . Grade 1 (early archaic Homo sapiens): included Bodo 1, Saldanha 1, Ndutu, Kabwe 1, and Eyasi. Grade 2 (late archaic Homo sapiens) : included Omo 2, Laetoli H 18, Florisbad , and possibly Kabwe 2 and Eli ye Springs 11693. Grade 3 ([early] anatomicall y modern Homo sapiens) included : Omo 1, the material from Klasies River Mouth and Border Cave, and possibly Singa. Generally , Brauer's placement of the hominids is close to that of Stringer et al. ( 1979), although the positions of some individuals vary. Brauer argued that Grade 2 hominids were contemporaneous with many of the members of Grade 3 (the direct dating of Florisbad placed it earlier than the Grade 3 material). As the chronolo gical placement of particula r and/or individuals has been refined, Brauer (Brauer et al. 1997) has revised the transition between the groups to 700500,000 years BP for Homo erect us/archaic Homo sapiens,

Other workers have also suggested that anatomically modem Homo sapiens originated quite early (in excess of 100,000 years BP) in south and/or east Africa , and have alluded to the possibility that this region was the source area for modern crania of a younger date than found in other areas (Beaumont 1980; Beaumont et al. 1978; Butzer et al. 1978; Day 1969; Singer and Wymer 1982; de Villiers 1973, 1976). For example, Singer and Wymer suggested that the material from Klasies River Mouth ...confirm not only that Homo sapiens sapi ens occupied this southern region of Africa by the

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350-250,000 years BP for early/late archaic Homo sapiens and at least 150,000 years BP for the origin of Homo sapiens. McBrearty and Brooks (2000) adopted a similar Grade model, but acknowledged that there were problems with the allocation of specimens to some of the Grades. Unlike Brauer, McBrearty and Brooks (2000) included late Homo erectus material along with early archaic Homo sapiens included Bodo 1, Saldanha 1, Ndutu and Kabwe in their Grade 1.

... intermediate, metrically and non-metrically, in an African series spanning about 600,000100,000 years ago ... (2003:745) Such an intermediate assessment of the Herto Bouri hominids is consistent with the overall morphology of the material. Herto 1 is less archaic than the Bodo, Saldanha and Kabwe hominids, but appears to share similarities in the angulation of the occipital and form of the occipital torus. Herto 1 is larger and more robust in the occipital and supraorbital regions than the material from Omo (Kibish), Florisbad, Laetoli, Eliye Springs and Klasies River Mouth. Herto 1 also links sub-Saharan Africa with north Africa as it has morphological parallels with the slightly younger Jebel Irhoud hominids, especially Jebel Irhoud 1. There are also morphological parallels between Beret ER-3884 and Jebel Irhoud 1, including thin vault bones, relatively high occipital angle, some lambdoidal protrusion and well-developed arched supraorbital torus displaying some lateral thinning. Whilst Herto 1 and Jebel Irhoud 1 display similarities of the face and vault, such as a relatively steep frontal, broad interorbital region, some occipital protrusion and a prominent occipital torus, a broad and flat face with squarish orbits, a broad nasal aperture, marked incisura malaris and alveolar prognathism, Jebel Irhoud 1 does have more prelambdoidal flattening, a smaller supraorbital region, shorter face and a taller nasal aperture.

Within the sub-Saharan African hominid sample evolutionary trends are evident (Brauer 1984; Rightmire 1989; Smith 1985; see also the results from the multivariate analyses). These include a decrease in overall robustness (especially of the face, and supraorbital region), a reduction in the size of the face, especially height, and facial prognathism, less postorbital constriction, greater elevation of the frontal squama, the separation of the superciliary arch from the supraorbital trigone, an increase in parietal length and degree of bossing, a decrease in mid-sagittal keeling, and changes in the form of the occipital squama. A gradation in the morphological features of the individual crania also illustrates an evolving lineage. The earlier Middle Pleistocene crania are, allowing for sexual dimorphism and chronological differences, morphologically similar to each other. Bodo 1 is more robust and displays a shorter flatter face with a greater degree of facial prognathism than Kabwe 1, but they are also morphologically similar. In many features, especially robustness and overall cranial form, Saldanha 1 resembles Kabwe 1, while in other features, such as occipital angulation and a laterally thinning occipital torus, it is closer to Ndutu. Both Saldanha 1 and Ndutu display some degree of parietal bossing and a lack of midsagittal keeling, which are features that are more commonly found on the later crania such as Omo 1, Laetoli H18, Florisbad, Herto 1, Border Cave 1 and Singa. Saldanha 1 is also similar to Eliye Springs 11693 in the sagittal curvature of the frontal bone, and in having a low position for maximum cranial breadth, something that is shared with Ndutu, Kabwe 1 and Omo 2. The Kabwe 2 maxilla displays similarities to the more robust, and probably earlier, Kabwe 1 cranium although it is smaller and has a shallow canine fossa. In these feature Kabwe 2 is similar to Laetoli H 18, Eliye Springs 11693, Florisbad and the earlier Ndutu cranium.

These morphological parallels document a general gradation in cranial form through time from the earlier robust and archaic hominids such as Bodo 1, Ndutu, Saldanha 1 and Kabwe 1, via intermediate forms such as Omo 2, Florisbad, Eliye Springs 1693, Eyasi 1, Herto 1 Beret ER-3884 and possibly Kabwe 2 which display, to varying degrees, similarities to the earlier and later material, through to more modern appearing crania like Laetoli H 18 and Singa, and finally to anatomically modern Homo sapiens that are found throughout sub-Saharan Africa by at least the terminal Pleistocene (Fig. 2:2). As McBrearty and Brooks observed In contrast to the pattern seen in western Europe, eastern Asia, and the Near East, there is a series of closely related taxa that provide plausible ancestors for H. sapiens, and the record shows no abrupt discontinuities. (2000:482-3) However, there are some disconformities in the sequence. The earlier robust and archaic hominids display morphological variation that may be related to sexual dimorphism and/or chronological differences with the smaller less robust and possibly female Ndutu cranium earlier than the probably male Kabwe 1. The intermediate forms such as Omo 2, Florisbad, Eliye Springs 1693, Eyasi 1, Herto 1 and Beret ER-3884 display both similarities as well as a wide range of morphologies. Beret ER-3884, if dated to 270.000 years BP, is less robust than later hominids such as Omo 2 and Herto 1. A more obvious disconforrnity is caused by the early dating of the essentially modem Omo 1 cranium and the Border Cave material and the more gracile remains from Klasies River Mouth, especially mandible KRM 41820, which Singer and Wymer claimed . .. represents the earliest Homo sapiens sapiens directly associated with an MSA culture in southern Africa or elsewhere. (1982:148)

Omo 2 displays a mosaic of morphological features that may be paralleled on both earlier and later crania. The occipital squama ·of Omo 2 recalls both Saldanha 1 and Kabwe 1, while its mid-sagittal keeling parallels Bodo 1 and Kabwe 1. The sagittal curvature of the parietals and the supraorbital region of Omo 2 are similar to those of Laetoli H18. As on Omo 2 and Laetoli H 18, the superciliary arch and supraorbital trigone are easily distinguished on Omo 1, Florisbad and Singa. Omo 2, Florisbad and Border Cave 1 share a broad frontal, while Laetoli H 18, Eli ye Springs 11693, Florisbad, Herto 1, Border Cave and Singa all have a steep supraglabellar portion of the frontal bone. The wide biorbital and interorbital regions of the Florisbad upper face approximate those of Herto 1, Eliye Springs 11693, Beret ER-3884, Laetoli H18, Kabwe 1 and Bodo 1. White et al. described the Herto Bouri material as

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poorly dated , to conclusively demonstrate human origins on the African continent. (1995:42)

As mentioned earlier, the Klasies River Mouth material has been used to demonstrate the appearance in sub-Saharan Africa of anatomically modem forms prior to 100,000 years BP (Beaumont 1980; Beaumont et al. 1978; Brauer 1984a, 1984b, 1984c, 1989a, 1989b; Butzer et al. 1978; de Villiers 1973, 1976), which parallels early dates for their appearance in the Levant. Prior to the revised dating of the material from western Asia, Beaumont stated These key data, [Klasies River Mouth material] complementing those from Border Cave, clearly demonstrate in view of the absence of contrary evidence from elsewhere, that the genesis of modern man took place somewhere south of the Sahara, and at a time certainly prior to 120-130,000 B.P . (1984:78) The maxillary fragment from Klasies River Mouth AA43 , however, fits within the general morphological gradation in that it displays some morphological similarities to archaic hominids such as those from Kabwe 1, although smaller in size, (Deacon 1989), and in overall size and proportions to Laetoli HI 8 (Rightmire and Deacon 1991). The Klasies River Mouth postcranial material, zygomatic KRM 16651 and robust mandibles KRM 13400 and KRM 21776 are also consistent with a morphological graduation from more archaic earlier material towards a more modern morphology .

Regional Features Both the Replacement Hypothesis and the Multiregional Hypothesis argue that anatomically modem humans evolved in situ in sub-Saharan Africa. Coupled with this view bas been the assumption that modern regional characters should have a relatively high antiquity (pre-dating the late Pleistocene) in this area (Wolpoff et al. 1984; Stringer and Andrews 1988: table 1, but see also Stringer and Brauer 1994). Rightmire (1984:32) . has suggested that if the datings for the hominids from Klasies River Mouth and Border Cave are correct "it can be stated that people very much like living populations (San or Negroes) have been resident in southern Africa for a long time ". This view has been adopted by other authors. Delson (1988:206) , for example , stated in a discussion of the origin of anatomically modem humans that "The southern African fossils are thought to show African regional features". Wolpoff ( 1989b:66) has also argued that the earliest modem hominid material from sub-Saharan Africa, whether it pre- or postdates 100,000 years BP, is "fundamentally African" in its morphological features. He suggested, therefore, that the hominids making up this material (Omo 1, and the Border Cave and Klasies River Mouth material) are the "earliest modem Africans, morphologically as well as geographically". Wolpoff goes on to state that ...they establish the presence of an African morphological complex in modems prior to their leaving Africa ... (1989b:66)

If the proposed early dates for Omo 1 and the hominid material from Border Cave and Klasies River Mouth were correct, they would have been contemporaneous with morphologically more archaic forms such as Laetoli H18 and Eliye Springs 11693, but as has been discussed in detail earlier, there are problems with the dating of these fossils. There are problems with the reliability of the technique used to date Omo 1, and all the cranial and mandibular Border Cave hominids are probably burials from higher levels than they were derived from and so much younger than the dates that are attributed to them.

However , a detailed list of morphological features that document "regional continuity" in sub-Saharan Africa has not been proposed, although a number of fossils (Saldanha, Florisbad, Border Cave, Klasies River Mouth, and Singa) have been said to display features that are present on modem Africans and so document morphological continuity in subSaharan Africa .

The situation at Klasies River Mouth is more complex because there are both robust and more gracile individuals present. The material is definitely from Middle Stone Age levels, but their placement within this period is not perfectly clear . For example, the most modem mandible , KRM 41820 from Shelter lB, may be quite a bit younger than the date given to it by the excavators (Binford 1984, 1986a, 1986b). The bulk of the hominid material is probably less than 100,000 years BP and there is the possibility that the earliest hominids from the site were robust and archaic , and that they were followed by more gracile and modem looking hominids (but see Deacon 1989). The evolutionary progression that may be represented at Klasies River Mouth parallels that which has occurred throughout the rest of south and east Africa. Deacon ( 1989; Deacon and Geleijnse 1988), however, contends that there was a single population with extreme sexual dimorphism present at Klasies River Mouth. At present , the suggested early Upper Pleistocene date for this material is not a secure enough base upon which to construct a parsimonious hypothesis claiming very early appearance of anatomically modem man in sub-Saharan Africa. As Mann has observed: An examination of the present fossil evidence documentin g the early appearance of modem human-like people s in sub-Saharan Africa suggests they are either too incomplete , or

What follows is an evaluation of the usefulness of the features that have been identified as demonstrating a close phylogenetic relationship between some Middle and early Late Pleistocene sub-Saharan African hominids and modem African groups. Saldanha The size and shape of the mandibular fragment from Saldanha , and reconstructions of it, suggest that the ramus was short and broad with a shallow sigmoid notch. Drennan and Singer (1955:365) felt that Saldanha 2 preserved the proportions of "Bushman , Hottentot and pre-Khoisan jaws" . However , a similar ramus form can be found on the Mauer mandible from Europe (Drennan and Singer 1955; Kraatz 1985a, 1985b ), while the almost complete mandible from Lake Baringo , Kenya , also has a broad ramus (Rightmire 1980).

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hominids. She found that Border Cave 1 was not "closely related to recent Khoisan or South African Negro populations", but did suggest that it displayed Negro features such as dolicho-orthocrany, absence of frontal narrowing, and an ovoid vault (de Villiers 1973:238). The dimensions of the Border Cave 2 mandible were found to fall within the range of South African Negro mandibles. De Villiers found Border Cave 3, the infant skeleton, to demonstrate ...a cluster of metrical and morphological features which align it with the South African Negro infant, as well as morphological features which relate it to the Khoisan infant. (1973:243) These Negroid similarities include limb proportions, and mandibular dimensions, while the Khoisan similarities are centred on the morphology of the mandible. In conclusion de Villiers, following Wells (1969), suggested that the Border Cave hominids ...may be representative of a proto-negriform, basic Homo sapiens afer population from which the Negro and Khoisan peoples diverged. (1973:244) The metrical and morphological features of Border Cave 5 are also said to favour a link with the South African Negro (Beaumont 1980; de Villiers 1976). In another multivariate study (discriminant analysis) de Villiers and Fatti (1982) found Border Cave 1 to fall within a Negro male distribution, and concluded that it had Negroid affinities.

Florisbad Wells (1969:172) suggested that the hominid from Florisbad was an "undifferentiated protonegro-protobushman" type, whereas Dreyer argued that ...the Florisbad skull belongs to the Pre-historic South African race - the Bushman, of which he is a very early and very primitive representative. (1936:23) Hoffman ( 1962:282) supported Dreyer's contention that the Florisbad hominid was "Bushmanoid", and "represents the ancestral stock of the Bushman race". He outlined features that are said to demonstrate the morphological link with the modem Bushman type. These features, based on Dreyer's initial reconstruction of Floris bad that was flawed (see earlier discussion and Clarke 1985; Rightmire 1978a), include obliquely angled rectangular orbits (now more subrectangular), lack of an ophryonic groove, prominent parietals, presence of a marked canine fossa (now known to be only slightly developed), flat nasal bones, small nasal aperture, and microdont teeth. However, these morphological parallels (some of which are now known not to be accurate) are very general, and could be used to suggest similarities with crania from other geographical regions outside of subSaharan Africa. For example, Galloway (1937:10) suggested that Florisbad displayed a combination of features that "recur in the Australian skull and in the Boskop physical type". Flat nasal bones are a suggested "regional feature" for east Asia (see Chapter 7; Weidenreich 1943b). Rightmire, after a reanalysis of the Florisbad hominid, concluded that Dreyer's attempts to tie the cranium into the ancestry of the Bushman may be viewed as extreme ... (1978a:484) and that Florisbad ...is not noticeably Bushman-like, and there is no evidence to support special ties with any one group of living Africans. (1978a:479)

A major problem with the analyses carried out by de Villiers is that it was assumed that the Border Cave hominids must have had a close morphological relationship with modem African groups, and so these were the only populations that the material was compared with. As de Villiers and Fatti stated ...the classification probabilities have been evaluated on the assumption that the unknown has, in fact, come from the populations considered ... (1982:323, 327) We do not know therefore, if the Border Cave material in general, and Border Cave 1 in particular, might not be more similar to modem populations from other geographical regions than they are to modem African groups. De Villiers and Fatti also contend that the classification probabilities of the modem populations that are not greater than 0.25 indicate that ...a fair amount of uncertainty still exists with regard to the true classification of the respective unknown. (1982:327) None of the classification probabilities for any of the Border Cave hominids, in relation to the individual modem populations, was greater than 0.25. This would mean therefore, that a great deal of "uncertainty" must surround any conclusions based on these results. It is only when the classification probabilities are combined and scaled so that there are only male and female Negro and San samples, that the Border Cave values get above 0.25. When this is done Border Cave 1 and 2 have a higher probability of belonging to the male Negro sample, whereas Border Cave 5 has a higher probability of belonging to the San male sample.

Along with the more general morphological similarities (relatively bulging frontal, broad face, wide interorbital region, and those mentioned by Hoffman [1962) and discussed above), Florisbad does display at least one specific Khoisan feature in that it has an interparietal groove. This feature, however, is also found on 33.6% of male and 37.8% of female South African crania studied by de Villiers (1968), although a wide interparietal groove like that on Florisbad, is only found on 3.4% of male and 8.8% of female crania. Florisbad also displays a number of general Negroid features (Brauer 1978; de Villiers 1968) including a frontal squama that is evenly curved in all directions, sub-rectangular orbits, wide interorbital region, and curved parietals, but these do not demonstrate any particular relationship because the features are commonly found on non-Negroid crania, including Khoisan material. Border Cave Wells ( 1969: 172) also suggested that Border Cave 1 was an "undifferentiated protonegro-protobushman" type. A number of attempts have been made to try and identify which modem African group the hominid material from Border cave is closest to. De Villiers (1973) carried out morphological and multivariate (discriminant analysis and Mahalanobis' Generalised Distance statistic) analyses of the Border Cave

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There are also theoretical problems in using discriminant analysis to allocate fossil hominids to modem populations (Campbell 1978; Corruccini 1975a; comments in Rightmire 1979). Corruccini contends that discriminant analysis was ...not designed to indicate relative affinities of unrelated individuals ...or to be applied to outside groups not included in the original computation ... (1975a:3) This is how de Villiers and Fatti (1982) used the method in the studies of the Border Cave hominids. To use discriminant analysis properly the individual that is being investigated must belong to one of the groups in the analysis. This is something that cannot be known with any certainty for fossil hominid material such as that from Border Cave.

From typicality probabilities Ambergen and Schaafsma (1984) found that Border Cave 1 was not "typical" ofany of the eight modem groups they compared it to. They state ...our computations suggest that Border Cave is certainly not a random drawing from any of the populations involved. (1984:130) Ambergen and Schaafsma (1984) go on to suggest that if it were assumed, even though they think it is doubtful, that Border Cave was a member of one of the modem African groups being used (Hottentots were not included), posterior probabilities indicate that it would be either the Bushman male or Zulu male groups. Using Mahalanobis' distances they found Border Cave 1 to be closest to Bushman males (Ambergen and Schaafsma 1984).

There are also problems in the way that de Villiers used Mahalanobis' Generalised Distance statistic . This statistic requires relatively large samples for the populations being compared, which does not seem to be the case in de Villiers' study. Corruccini also stated that ...it would be inappropriate to compare , say, a single individual with a population using D2 ... (1975a:3) This is exactly how de Villiers has utilised the statistic in her examination of Border Cave 1.

Campbell (1984) used individual squared Mahalanobis' distances and typicality and group membership probabilities. He found that with equal covariance matrices Border Cave 1 was most closely related to Khoisan groups than to Negro groups, and that with unequal covariance matrices Border Cave 1 was most similar to Bushman males. Campbell (1984) concluded that the typicality probabilities support Rightmire's (1979, 1981c) conclusion that Border Cave 1 was Bushman-like, although it is noted that if the typicality calculations were based on the chi-squared distribution instead of a Gaussian distribution Border Cave 1 would have been "atypical" for all the modem African groups considered.

Rightmire has also conducted a multiple discriminant analysis so as to ...explore the affinities of Border Cave with eight groups of modem Bushman, Hottentot, and South African Negro crania ... (1979:24) Rightmire ( 1979:25) found that Border Cave was "well within the range of modem African variation for the measurements used", and that it lay "closest to the Hottentot centroid". These results are in direct contrast to those of de Villiers and Fatti (1982). Many of the problems that have been outlined in relation to the analyses of de Villiers (de Villiers 1973, 1976; de Villiers and Fatti 1982) can also be levelled at Rightmire (cf. the comments that accompany Rightmire's 1979 paper, especially those by C.P. Groves and S.R. Wilson and Corruccini 1992). Campbell (1980) has also criticised Rightmire's ( 1979) use of multivariate techniques. So as to answer Campbell's ( 1980) criticisms, Rightmire ( 1981c) reanalysed his data and considered total Mahalanobis Generalised Distance instead of only the first six axes as in his previous study (Rightmire 1979). Border Cave was again found to be closest to Bushman and Hottentot male centroids (the results were the same when two "suspect;' measurements were left out and only modem male samples were used). Rightmire (1981c) did observe that Border Cave was relatively far from all of the modem groups. Rightmire concluded that ...there remains a substantial body of metric evidence to support allocation of the BC-1 cranium to a large Bushman-like population ... (1981c:200)

Van Yark, Bilsborough and Dijkema (1989) used a number of multivariate statistics, including Mahalanobis' distance, to compare measurements of Border Cave 1 taken by Howells and Rightmire with both recent African and non-African population samples. They found that Border Cave 1 did not fit with any of the samples compared and that it was not definitely more like recent African populations than it was like other recent populations. In fact they found a general preference for Australian populations (van Yark et al. 1989). Overall, however, Border Cave 1 was found to be significantly different from all recent population samples (van Yark et al. 1989). Corruccini (1992) reanalysed Rightmire's (1979) material and found that Border Cave 1 fell outside the "morphological envelop" of the modem African sample and was a "classic statistical outlier" Another major problem with any multivariate analysis of metrical data from Border Cave 1 is that there are few measurements that can be established with any certainty, and so one must either use very few measurements or measurements that are estimates. For example, Rightmire (1979, 1981c) used only 11 or 9 measurements, while de Villiers (1973) used eight measurements, four of which were regarded as questionable . Border Cave 1 does display morphological features that are typical of both Khoisan crania (relatively high bulging frontal, wide interorbital region, small mastoids, and prominent mastoid and supramastoid crests) and Negroid crania ( frontal evenly curved in all directions, no ophryonic groove, slight glabella development, little frontal narrowing, a wide interorbital region, a moderately deep digastric fossa, and a small mastoid process associated with a large mastoid crest), but these are again very general morphological traits which do not, by themselves , demonstrate any close

Ambergen and Schaafsma (1984) and Campbell (1984) have also used multivariate techniques to evaluate the relationship of Border Cave 1 to modem African groups.

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None of the Middle and/or early Upper Pleistocene hominid material from sub-Saharan Africa (except possibly Florisbad with its interparietal groove) can be shown to display "regional features" that document a closer phylogenetic relationship to modem African populations than to groups from other geographical regions. Crania that are clearly linked to modem African populations do not appear until the terminal Pleistocene (Brauer 1978; Brothwell 1963; Rightmire 1975, 1978; Singer 1958; Wolpoff 1980a).

phylogenetic relationships. Border Cave 5 also has a short and broad ramus, which is commonly found on Hottentot and Negro mandibles. As mentioned in the discussion of Saldanha 2, however, a similar ramus form can be found on fossils from outside of sub-Saharan Africa, including the Mauer mandible from Europe. Klasies River Mouth Some of the Klasies River Mouth hominid sample are said to display modern African morphological features. Singer and Wymer stated The features in the isolated KRM specimens, and especially 16424, would raise the possibility of a mosaic or pattern of generalized negroid features ...that would indicate ancestral forms of modem Negro, Bushman, and Hottentot peoples. (1982: 148) The Cave 1 frontal fragment, KRM 16425, is said to have the flat nasal root and limited development of the supraorbital region of many modem African crania (Rightmire 1978b; Singer and Wymer 1982), although it has been suggested that this fragment is from a juvenile (Frayer et al. 1993). The Shelter IA parietal fragment, KRM 41658, is suggested to indicate a low vault like that of the Bushman, and the narrow vault of the Hottentots and Negroes (Singer and Wymer 1982; but see Rightmire and Deacon 1991). The Shelter 1B mandible, KRM 41815, has a squarish ramus resembling Hottentot and Negro mandibles, whereas the short corpus is said to indicate a small broad Bushman-like face (Singer and Wymer 1982). The estimated size of the teeth of this individual falls within the size range of modem Hottentots and Negroes (Singer and Wymer 1982). Mandible KRM 16424 has a low ramal height, small five-cusped molars and is mesotaurodont, all of which are said to be Bushman features (Singer and Wymer 1982). Singer and Wymer (1982:148) stated that "in MSA I times a very sapient individual was present at KRM, with a broad and possibly orthognathous face, with a mandibular ramus somewhat in between that of a Bushman and a Negro (in modem terms), and with teeth possibly in a like size category".

Conclusion The results of the multivariate analyses and morphological comparisons indicate that there is morphological continuity within the corpus of late Middle and early Upper Pleistocene hominid crania from sub-Saharan Africa. They also suggest that there was an in situ development of anatomically modem forms in this region sometime after 100,000 years BP and before the advent of the Late Stone Age at around 40,000 years BP (Deacon 1984). The archaeological record also supports these findings in that it documents ...an unbroken sequence of evolution in artefact-making from the Earlier Acheulean to the Later Acheulean and then a change eliminating the heavy bifaced Earlier Stone Age artefacts but placing new emphasis on the ancient unifacial flake component at the beginning of the Middle Stone Age. (Mason 1962: 156) Klein also stated that Acheulean assemblages ...changed through time ...but the pace of change appears to have been agonizingly slow. (1983:31) while Volman argued that The earliest MSA assemblages ...do not show technological improvements over ESA ones ... (1984:218) and that ...the only widespread, convenient marker for distinguishing ESA from MSA seems to be the presence or absence of large cutting tools. (1984:192)

However, all of the morphological parallels between the Klasies River Mouth hominids and modem African groups that have been discussed are very general, and so do not necessarily demonstrate any close phylogenetic relationships. An example of the general nature of the morphological parallels is evident in the way that the small teeth of KRM 16424 fall outside the range for pygmies and Bushmen, but within the range of Europeans (Singer and Wymer 1982). Does tooth size therefore indicate a closer phylogenetic relationship between KRM 16424 and Europeans than between KRM 16424 and modem Africans?

As these statements suggest, there is strong evidence to demonstrated the slow in situ development of Upper Acheulean assemblages and a transition into Middle Stone Age assemblages (Clark et al. 2003; Volman 1984). The timing of the transition may vary in different regions, but there is no evidence to suggest a truncation of the developmental sequence. Although Middle Stone Age assemblages vary more in time and space than Early Stone Age assemblages, there is still a high degree of continuity within it (McBrearty and Brooks 2000; Volman 1984). It has been suggested that the Howieson's Poort component of the MSA was intrusive. Singer and Wymer concluded that There is certainly nothing to suggest that the Howieson's Poort stage at KRM evolved from the preceding MSA industry ... (1982:209) and that At some sites, such as KRM and the type site, the industry is in such marked contrast to the

Singa The Singa calvaria has often been regarded as having protoBushman or Bushrnanoid affinities, especially the posterior of the vault (Briggs 1955; Wells 1972; Woodward 1938). However, it has been suggested that the hominid may display pathological alteration (Brothwell 1974; C.B. Stringer ~ers. comm.), which can account for its resemblance to Kh01san crania. The extreme position of the Singa calvaria on the correspondence analysis (Figure 2: 10) would support this proposition.

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traditional MSA that quite populations might be considered.

separate

earlier trends and traditions that were enhanced and supplemented (McBrearty and Brooks 2000).

(1982:206) However , sites such as Border Cave demonstrate that the Howieson's Poort was an integral part of the MSA culture (and stratigraphy) during which some new ideas, techniques, and/or stylistic trends were adopted, and that it was not an intrusive and entirely separate industrial complex (Beaumont 1984; Parkington 1990; McBrearty and Brooks 2000; Volman 1984). However, it is worth noting that the origin of anatomically modem Homo sapiens in sub-Saharan Africa was not accompanied by an "archaeological revolution" , but rather a continuation of

There is then, strong evidence to support an in situ developmental/evolutionary sequence within the subSaharan African region of both skeletal and artefactual material. As argued by McBrearty and Brooks ... both human anatomy and human behaviour were intermittently transformed from an archaic to a more modem pattern over a period of more than 200,000 years. (2000:458)

60

CHAPTER THREE. NORTH AFRICA The pathological changes that appear to have affected Sale may account for its extreme position in a principal components analysis of the parietals carried out by Brauer (1984). It is worth noting that the Singa calvaria, which also appears to have been affected by pathological changes (see discussion in Chapter Two), is also extreme on this analysis (Brauer 1984a).

Introduction A large number of hominid remains have been recovered from north Africa, especially in the northwestern comer (Fig. 3:1, Table 3:1). Most of this material appears to be either older or younger than the period being dealt with, and so it will not be discussed in any detail. The dating of much of this material however, is not conclusive and so chronological comparisons to other regions are preliminary until greater clarity in dating is achieved ( see McBrearty and Brooks 2000; Hublin 1985 for a discussion of the dating of the north African material). There is a large collection of anatomically modem late Upper Palaeolithic skeletal material from the region, particularly from the sites of Afalou-bou-Rhummel in Algeria and Taforalt in Morocco, and earlier fragmentary hominid remains, especially mandibular, from Ternifine (now Tighenif), Thomas Quarries and Sidi Abderrahman (Fig. 3: 1; Oakley et al. 1977). This later material, which appears to be older than 400,000 years BP (Hublin 1985; McBrearty and Brooks 2000; Schwarcz 1982; cf. Jaeger ~975, 1981; Saban 1975, 1977 for younger dating schemes), 1s essentially Homo erectus in overall morphological configuration (Howell 1978; Oakley et al 1977). It is worth noting that Schwartz and Tattersall (2000; see also Dobson and Trinkaus 1990) have identified features on the symphyseal region of the Ternifine (Tighenif) mandibles such as a central keel, that typify extant Homo sapiens. McBrearty and Brooks (2000) placed most of this material, along with the Sale, Bodo, Ndutu, Kabwe, Saldanha and subSaharan African Homo erectus material into their Group 1.

If the morphological features of the posterior of the vault are dismissed due to pathological changes, Sale has been regarded as an evolved Homo erectus (see Day 1986), but if the overall morphological pattern is considered and the "extreme features" of the occipital disregarded, Sale can be regarded as a small (possibly female) archaic Homo sapiens (see Hublin 1985). The Sale calvaria is currently date to around 400,000 years BP based on linear uptake electron spin resonance ages of 389-455,000 years BP (quoted in McBrearty and Brooks 2000). McBrearty and Brooks (2000) placed Sale, along with the material from Ternifine (Tighenif), Sidi Abderrahman, Bodo, Ndutu, Kabwe, Saldanha and sub-Saharan African Homo erectus material into their Group 1. It is worth noting that if Sale is dated to only 400,000 years BP and is classified as an evolved Homo erectus, it would be more recent than or contemporary with sub-Saharan African material such as Bodo and Ndutu, that is referred to as archaic Homo sapiens. It would however, be contemporary with Homo erectus material from China (Chapter 7).

Mifsud-Giudice Quarry, Kebibat - Rabat Sale

The fragmentary remains of an adolescent from MifsudGiudice Quarry, Kebibat, near Rabat, Morocco, also appear to have belonged to either an evolved Homo erectus or archaic Homo sapiens (Fig. 3:1; Brauer 1984a; Howell 1978; Saban 1975, 1977; Schwartz and Tattersall 2000; Wolpoff 1980a). These remains appear to be more recent than the Sale calvaria, but are older than 200,000 years BP (Hublin 1985; Steams and Thurber 1965). The maxilla lacks a canine fossa and indicates pronounced alveolar prognathism, while the mandibular fragment has a robust body with an almost vertical anterior symphysis lacking a chin (Schwartz and Tattersall 2000). The well-rounded occipital fragment displays a high occiput with a reduced nuchal torus, along with an archaic middle meningeal vein pattern (Brauer 1984a; Howell 1978; Saban 1975, 1977).

A calvaria, along with fauna! material but no stone tools, was recovered from an open quarry near Sale, Morocco (Fig 3: 1; Day 1986; Hublin 1985; Jaeger 1975, 1981). The Sale calvaria displays a combination of erectine and more modem features (Hublin 1985; Jaeger 1975, 1981; Laitman and Cretin 1980). It is relatively long and low with marked postorbital constriction, frontal keeling a small cranial capacity and lacks an angular torus. Sale also has a number of modem appearing features on the posterior portion of the vault such as the form of the cranial base, the d~velopment of the parietal bosses, the rounded occipital with a reduced nuchal plane and a weak occipital torus. However, some of these modem features appear to be the result of pathological changes to the posterior of the vault (Hublin 1985; McBrearty and Brooks 2000; Stringer et al. 1984; Smith 1985). Hublin (1985) indicated that there was an irregular thickening of bone at the level of the linea nuchae inferior as a result of an expansion of the diploe, and not of the inner or outer tables as in Homo erectus (Kennedy 1985).

Jebel Irhoud-Djebel Ighoud · Four hominid fossils have been recovered from a barytes quarry on the northeastern side of the Jebel Irhoud, south of Safi, Morocco (Fig. 3: 1; Ennouchi 1962, 1968, 1969; Hublin et al. 1987; Hublin and Tillier 1981).

Clarke ( 1990) has identified similarities between the Sale calvaria and the Ndutu specimen including overall size, outline from above and behind (i.e. degree of postorbital constriction and parietal form) and morphology of the supramastoid region.

Jebel Irhoud 1 (Plate 3; Brauer 1984a; Ennouchi 1962; Howell 1978; Wolpoff 1980a) consists of an adult cranium that is relatively long, low and broad, with little postorbital 61

Dar-es-Solt an Jebel

.

Rabat . sate~ lrhoud •\emara

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Figure 3:1. Map showing North African sites mentioned in text.

Hominid Dar-es-Soltan 5 El-'Aliya Haua Fteah material Jebel Irhoud 1 Jebel Irhoud 2 Jebel Irhoud 3 Jebel Irhoud 4 Jebel Saharba Rabat Sale Sidi Abderrahman Temara Ternifine material Thomas Quarries material Wadi Halfa

Orhdnal Or Cast Published Cast Casts Cast Cast Published Published Published Original Original Cast Cast Casts Casts Published

Abbreviations

--

--Jll

-----

Principal Source Of Data Ferembach 1976b Cast Casts Cast Cast Hublin and Tillier 1981 Hublin, Tillier, and Tixier 1987 Anderson 1968 Original Original Cast Cast Casts Casts Greene & Annelagos 1972

Table 3:1. North African hominids examined, abbreviations for Figures and Tables, and source of data.

although there is a centrally prominent torus that tapers laterally and has sulci above and below . There is marked pre-lambdoidal flattening . The temporal squama are long and low. The tapering mastoid processes are quite prominent. The supraorbital torus is prominent , but not

constriction. The supraglabellar portion of the frontal squama is quite steep, while the remainder recedes gently . The parietal s are relatively long and flat, with only slight bossing. The rounded occipital protrudes , but does not display a Neanderthal configuration (Santa Luca 1978),

62

Three mammal teeth derived from a level immediately overlying Jebel Irhoud 4 have been used for electron spin resonance (ESR) dating (Griin and Stringer 1991). The age estimates ranged between 90-125,000 years BP for early uptake (EU) and 105-190,000 years BP for linear uptake (LU)(Griin and Stringer 1991). The large age ranges for both EU and LU may indicate that the Jebel Irhoud site had a long depositional history. Griin and Stringer (1991) suggested that if the hominids were all recovered from near the cave floor and so low in the stratigraphic sequence, they could date to oxygen isotope stage 6, between 130-190,000 years BP (Table 5:1).

especially large, with the superciliary arches slightly more prominent than the supraorbital trigones. There is a marked ophryonic groove separating the supraorbital region from the rest of the frontal squama. The gracile face of Jebel Irhoud 1 is short, broad and flat, with large, low squarish orbits. The interorbital region is broad, while the nasal cavity is broad but short. The delicately constructed malars form a sharp angle with the maxillae, have a marked incisura malaris and display slight canine fossae. The face is orthognathic except for the alveolar region which projects markedly. The palate is broad and deep but not long, and the remaining tooth roots are large and robust.

The faunal remains from Jebel Irhoud and the LevalloisoMousterian artefacts, which are similar to those from other sites in the region including Haua Fteah, suggest a minimum age of between 40,000 and 60,000 years BP (Ennouchi 1962, 1963), but the site could be correlated with oxygen isotope stage 5 which would mean an age of between 80,000 and 120,000 years BP (Table 5:1; Griin and Stringer 1991; Ruhlin et al. 1987; Smith et al. 1989). This age would be consistent with the EU ESR dates obtained by Griin and Stringer (1991).

Jebel Irhoud 2 (Brauer 1984a; Ennouchi 1968; Howell 1978; Wolpoff 1980a) is an adult calvaria that, in many respects, resembles the Jebel Irhoud 1 cranium. The vault is relatively long, low and broad, with little postorbital constriction, long, flat parietals, long, low temporals, pre-lambdoidal flattening, and a protruding occipital. On Jebel Irhoud 2 the frontal, especially the supraglabellar region, is higher, the parietal bossing is more developed, the occipital is more protruding and angled, the nuchal plane is larger and the mastoid region is more prominent than on J ebel Irhoud 1. The prominent supraorbital torus of Jebel Irhoud 2 is more arched than that of Jebel Irhoud 1, and the superciliary arch can be more easily differentiated from the supraorbital trigone. The differences between Jebel Irhoud 1 and 2 may be due to sexual dimorphism, with the former being female and the latter male. Jebel Irhoud 2 is also said to have a patent metopic suture (Tobias 1968).

In the analyses Jebel Irhoud 1 took up one of two positions. It could be a late joiner to the European Neanderthal group (Figs. 3:2, 3:3, Table 3:2), where there were high loadings on the variables that describe the shape of the posterior cranium and the breadth of the lower face (Fig. 3:3, Table A:5). In another analysis Jebel Irhoud 1 grouped with Kabwe 1, and then joined with Steinheim, Gibraltar 1, Skhul 5 and Jebel Qafzeh 6 (Fig. 4:3, Table 4:2). There were high loadings on the variables that document cranial shape and facial size (Fig. 4:3, Table A:8). In the ordered similarity matrix Jebel Irhoud 1 was identified as being similar to both Neanderthal and non-Neanderthal material from Western Asia (Table 4:7). In earlier analyses Jebel Irhoud 1 also grouped with Kabwe 1 (Habgood 1982, 1984b; Habgood and Walker 1986).

Jebel Irhoud 3 (Ennouchi 1969; Ruhlin and Tillier 1981) is an infant mandible containing both deciduous and permanent teeth that was found at a lower level than the two adult crania, but in the same Levalloiso-Mousterian layer (Ennouchi 1969). The teeth are very large, and although there is not a true mental trigone all four components of it are recognizable on the nearly vertical symphysis. Schwartz and Tattersall (2000) suggest that there may be a central keel on the symphysis that expands inferiorly into a thickened inferior margin. The preserved left ascending ramus is relatively broad, vertically disposed, with a high coronoid process and slightly obtuse gonial angle. Jebel Irhoud 4 is a strongly built humerus of a child (Ruhlin et al. 1987).

Table 3:2. K-means cluster analysis with Jebel Irhoud 1. Analysis includes 72.6% of within-group variance. Group 1 Combe Capelle, Chancelade, Dolni Vestonice 3, Oberkassel 1, Oberkassel 2.

Establishing a date for the Jebel Irhoud material has proven difficult. Day ( 1977, 1986) suggests that Ennouchi felt that the fauna and stone tools demonstrated that the site was indisputably Middle Pleistocene in age. Biberson (1964) placed the site within the Ouljian marine transgression between the Presoltanian and the Soltanian (Jaeger 1975, 1981), while Oakley ( 1966) equated it with the Soltanian.

Group 2 Predmost 3, Predmost 4, Cro-Magnon 1, Mladec 1, Abri Pataud 1.

A radiocarbon date of greater than 32,000 years BP (Ny-73) was obtained from the upper part of the deposit (Coppens et al. 1968; Stringer and Burleigh 1981). However, Briggs ( 1968) stated that the radiocarbon dating at the site indicated an age of 32,000 years BP, and Oakley et al. ( 1977) gave the date as greater than 30,000 years BP. Ennouchi (1968) concluded that the deposits were beyond the effective range of the radiocarbon technique used.

Jebel Irhoud 2 was a late joiner to a group that included Neanderthal, archaic Homo sapiens and archaic modem crania. There were high loadings on the variables describing the breadth of the vault and the shape of the parietals.

Group 3 Steinheim, Kabwe 1, Skhul 5, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Jebel Irhoud 1.

The positions of the Jebel Irhoud crania are generally consistent with their respective morphologies. As is indicated by the results of the various analyses, Jebel Irhoud 1 does resemble the Neanderthals in the shape of its posterior 63

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Figure 3:3. Correspondence analysis with Jebel Irhoud 1. 64

cranium, with flat parietals, pre-lambdoidal flattening and occipital protrusion. The facial region of Jebel Irhoud 1 does not generally resemble that of Neanderthals. However, the large breadth of the nasal aperture of Jebel Irhoud 1 is reminiscent of the Neanderthals, although the height is much reduced in the north African hominid.

tripartite occipital torus with a suprainiac fossa (Santa Luca 1978; Table 5:6). Jebel Irhoud 1 does resemble Kabwe 1 in that they both have very broad faces, pre-lambdoidal flattening and occipital protrusion. In other respects, such as overall robustness or the degree of facial projection, they are quite different. There are also similarities between Jebel Irhoud 1 and some of the other sub-Saharan African crania such as Florisbad and Laetoli H 18, which also have steepish supraglabellar regions and moderately developed canine fossae. Wolpoff (1980a) has suggested that there are similarities between the two crania from Jebel Irhoud and the two Omo crania, although this similarity is limited with Omo 1 possessing a more rounded and less robust cranium and Omo 2 being more archaic than the Jebel Irhoud material. Jebel Irhoud 1 has a degree of facial prognathism that could match the Haua Fteah mandibles. The steep supraglabellar portion of the frontal bone of Jebel Irhoud 1 recalls that of Zuttiyeh, Tabun Cl, and Skhul 4 and 9.

Corruccini (1974a) found Jebel Irhoud 1 to be similar to the Neanderthals and Jebel Irhoud 2 to be more advanced, whereas in a later study (Corruccini 1992) Jebel Irhoud 1 and 2 were separated from the Neanderthals and grouped with Jebel Qafzeh 6, Shkiil 4 and 9, Omo 2 and Florisbad. Stringer (1974b) found Jebel Irhoud 1 to be closest to Saccopastore 1, Amud 1 and Skhul 5, and suggested that it was a generalised Neanderthal form. Stringer ( 1978) found that in a Pemose size and shape analysis of the neurocranium Jebel Irhoud was similar to the Neanderthal reference group, whereas in the analysis of the entire cranium Jebel Irhoud 1 was further separated from the Neanderthal reference group. Stringer (1978) found that Jebel Irhoud 1 did not have a Neanderthal face. Howells (1989) found Jebel Irhoud 1 to be grouped with Kabwe 1 and Skhul 5 and separated from "classic" Neanderthals from Europe.

There appears to be close morphological parallels between Jebel Irhoud 1, and to a lesser extend Jebel Irhoud 2, and the possibly 160,000 years old Herto 1 individual. Herto 1 also has a relatively steep frontal, broad interorbital region, prominent mastoid processes, some occipital protrusion and a prominent occipital torus. Jebel Irhoud 1 does appear to have relatively longer and flatter parietals, lower temporal squama and more pre-lambdoidal flattening than Herto 1. Whilst Jebel Irhoud 1 has a smaller supraorbital region, a shorter face and a taller nasal aperture, it shares with Herto 1 a broad and flat face with squarish orbits, a broad nasal aperture, marked incisura malaris and alveolar prognathism.

On a principal components analysis of the frontal squama, Brauer (1984a) found Jebel Irhoud 1 and 2 to be well separated. Although not positioned relatively close to other hominids, Jebel Irhoud 1 was found to be on the same half of the plot as Saldanha 1, Kabwe 1 and Laetoli H18. Jebel Irhoud 2 was on the same half of the plot as Omo 1, Florisbad, Border Cave 1 and Singa, and was positioned near anatomically modem African material. In a principal components analysis of the parietals, Brauer ( 1984a) found Jebel lrhoud 1 to be positioned on the archaic half of the plot. It was separated from the other archaic hominids, with Omo being closest to it. In an analysis of the face, Brauer (1984a) found Jebel Irhoud 1 to be positioned in an intermediate position between the archaic Petralona, Bodo 1 and Kabwe 1 and anatomically modem north African hominids.

In facial morphology Jebel Irhoud 1 also resembles east Asia material with its low squarish orbits and short, broad and flat face with delicately constructed malars that form a sharp angle with the maxillae and have marked incisura malaris (Pope 1991, 1992).

Simmons (1991) undertook a multivariate analysis of the frontal and supraorbital region and found strong links between the Jebel Irhoud hominids and material from the Levant and especially with some of the Jebel Qafzeh crania. Simmons ( 1991) did not find any close similarity with archaic or early anatomically modem humans from Europe.

Jebel Irhoud 2 also has a number of similarities with the Neanderthals, such as the protrusion of its occipital, but less so than Jebel Irhoud 1. The infant mandible, Jebel Irhoud 3, also lacks the typical Neanderthal morphology and is most similar to material from Skhul and Qafzeh (Hublin and Tillier 1981).

Ennouchi (1962, 1963), Tobias (1968) and Howell (1978) concluded that Jebel Irhoud 1 was a north African Neanderthal. Tobias suggested that Possibly, the Jebel lghoud remains represent the extreme westerly limit of the Afro-Asian Neandertals of the Upper Pleistocene. (1968:187)

Stringer et al. (1979) placed Jebel Irhoud in their Homo sapiens Grade 3, while Brauer (1984a) allocated Jebel Irhoud to his Homo sapiens Grade 2a (Neandertaloid Homo sapiens). McBrearty and Brooks (2000) placed the Jebel Irhoud material, along with Omo 2, Laetoli H 18, Haua Fteah and Mugharet el-'Aliya into their Group 2.

Although Jebel Irhoud 1 did, on occasions, group with the Neanderthals, the similarities are predominantly superficial and, as mentioned above, are confined to the shape of the posterior vault. Jebel Irhoud 1 does not posses the typical Neanderthal facial or occipital morphological pattern in that it has canine fossae, square orbits and subnasal prognathism (not mid-facial prognathism), a prominent but not especially large supraorbital torus, large occipitomastoid crests and a

In overall morphological configuration the Jebel Irhoud hominids resemble archaic Homo sapiens with some superficial resemblances to the Neanderthals. Stringer (1989b, pers. comm.) has argued that Jebel Irhoud 2 displays the closest approximation to a transitional morphology between archaic and modem humans of any crania that he has studied. As Briggs stated The appearance of both specimens [Jebel Irhoud 1 and 2] is indeed reminiscent of that of 65

the classic Neanderthal skull of La Chapelleaux-Saints but the resemblance is only superficial, a sort of optical illusion which fades away under careful examination. (1968:381)

As the comparisons made by Tobias (1967) demonstrate , most of the features present on the Haua Fteah mandibles are quite variable in their distribution on hominid mandibles and so are of limited use for classification purposes . The Haua Fteah mandibles do not, however , display the typical Neanderthal retromolar gap, horizontal-oval mandibular foramen pattern , taurodont molars or the posteriorly angled ramus with a shallow sigmoid notch with its deepest portion towards the condyle that is smaller than the coronoid process .

Haua Fteah Two mandibular fragments were recovered from LevalloisoMousterian levels within the cave of Haua Fteah on the northeastern coast of Libya (Fig . 3: 1; McBumey et al. 1953; Trevor and Wells 1967). A radiocarbon date of 47,000+3,200-2,300 years BP ("bone" fraction ; GrN-2023) and 40,700+/-1500 years BP ("rest" fraction ; GrN-2022) was obtained from charred bone in the same layer as the hominid remains (Vogel and Waterbolk 1963). Fluorine and nitrogen analysis has confirmed the association of the mandibular fragments and animal bone s from the same level (Oakley 1967; Oakley et al. 1977). McBr earty and Brooks (2000) cite preferred age estimates of >90,000 years BP and > 130,000 years BP for the Haua Fteah material.

Brauer (1984a) placed Haua Fteah in his Homo sapiens Grade 2a (Neandertaloid Homo sapiens). The Haua Fteah mandibles do not belong to Neanderthals and were most probabl y associated with crania similar to Jebel Irhoud 1. McBrearty and Brooks (2000) placed the Haua Fteah material , along with Omo 2, Laetoli H18, Jebel Irhoud and Mugharet el-'Aliya into their Group 2.

Mugharet El-'Aliya-Tangier (High Cave) The remains of two individuals were recovered from the High Cave , Mugharet el-'Aliya, near Tangier, Morocco (Fig. 3:1; Briggs 1955; Howe 1967; Senyurek 1940). El-'Aliya 1 is a fragment of a child's left maxilla with erupted and unerupted teeth, while el-Aliya 2 is a large and worn adult left second molar. El-'Aliya 1 is quite thick and has a horizontal nasal floor with a rounded inferior margin, blunt nasal spine , a bulging sub-orbital surface with no indication of a canine fossa, subnasal guttering, posterolaterally sloping zygomatic process and large teeth (Coon 1962; Howe 1967; Howell 1978; Senyurek 1940).

The young adult (given an age of 18-25 years) Haua Fteah 1 comprises the posterior portion of the left mandibular corpus with the worn second molar and the relatively unworn third molar in place and much of the left ascending ramus (McBumey et al. 1953; Tobias 1967). It has a relatively low and broad ramus, with short and blunt coronoid process and a shallow sigmoid notch with its deepest portion towards the midpoint. The anterior border of the ramus overlaps the third molar and so there is no retromolar gap. The second molar, which has a cusp pattern intermediate between the "Y5" and "+5" pattern , is larger than the third molar, which has a "+5" cusp pattern . The molars do not appear to be taurodont (McBurney et al. 1953). The mandibular foramen pattern is not of the horizontal-oval form (Tobias 1967).

The exact stratigraphic provenance of el-'Aliya 1 is unknown (Coon 1962), but fluorine and nitrogen analyses suggest that it derived from Layer 5 (Howe 1967; Oakley et al. 1977), a diminished or Final Aterian level. This level is equated with an equivalent level at Dar-es-Soltan that has a radiocarbon date of greater than 27,000 years BP ([UCLA-678B] Oakley et al. 1977). The stratigraphic position of el-'Aliya 2 remains in some doubt (Briggs 1968). McBrearty and Brooks (2000) cite a preferred age estimate of 60-90,000 years BP for the Mugharet el-'Aliya material based on dates for Aterian material elsewhere in north Africa .

The juvenile (given a most likely age of 12-14 years) Haua Fteah 2 consists of the left ascending ramus with the fully formed third molar crown present within its follicle (Tobias 1967; Trevor and Wells 1967). The rami is relatively low and broad , with a shallow sigmoid notch with its deepest portion towards the midpoint and a short pointed coronoid process. The third molar crown displays the "+5" pattern . Trevor and Wells (McBurney et al. 1953; Trevor and Wells 1967) and Howell ( 1978) suggested that the Haua Fteah mandibles had similarities with material from Tabfin, Amud and Shanidar, and so have Neanderthal affinities. Tobias ( 1967) found the Haua Fteah material to display similarities with the western Asian Neanderthals from Tabfin and Shanidar, the more modem material from Skhiil and Ksar 'Akil, and both earlier and later hominid material from Africa. He felt that .... the individuals represented by the Haua Fteah mandibles seem to have belonged to a widespread Afro-Asian neandertaloid population which had at least some of its genetic roots in the earlier hominid populations of Africa. (1968:185)

El-'Aliya 1 is of interest because it could only be, say, 30,000 years old (it could , of course, be much older), yet it has morphology that is reminiscent of the much earlier Rabat maxilla, which also lacks a canine fossa (Briggs 1955, 1968; Coon 1962), and contrasts markedly with the possibly chronologically closer Jebel Irhoud 1 face which is more gracile and has canine fossae. Brauer (1984a) placed Mugharet el-'Aliya in his Homo sapiens Grade 2a (Neandertaloid Homo sapiens). Senyurek (1940) and Howell (1978) have suggested that el-'Aliya 1 displays Neanderthal affinities, especially the lack of a canine fossa. However, all of its morphological features, including the lack of a canine fossa, may be found on earlier hominids from north Africa (cf. Coon 1962), which are not Neanderthals . McBrearty and Brooks (2000) placed the Mugharet el-'Aliya, along with Omo 2, Laetoli H18, Haua Fteah and Jebel Irhoud into their Group 2.

66

A more secure date for this material, especially Temara 1, is needed so as to successfully place it within an evolutionary framework.

Temara Fragmentary hominid remains have been recovered from Contrebandiers Cave, southwest of Rabat, Morocco (Fig. 3:1; Ferembach 1976a; Roche and Texier 1976; Vallois and Roche 1958).

Dar-Es-Soltan( e)

Temara 1, a mandibular fragment (Brauer 1984a; Howell 1978; Ferembach 1976a; Vallois and Roche 1958), has a low and robust body, a broad and low ramus with a shallow sigmoid notch, no retromalar gap, a relatively vertical symphysis with the beginning of a mental trigone and large teeth. It is said to display many similarities to the mandible from Rabat (Brauer 1984a; Howells 1978), although the symphysial region displays more fullness (Tobias 1968). Tobias (1971) suggested that there were also similarities between Temara 1 and mandibular material from the Cave of Hearths, South Africa, and Dire-Dawa, Ethiopia.

The remains of at least three hominids have been recovered from Dar-es-Soltan near Rabat, Morocco (Fig. 3:1; Brauer 1984a; Debenath et al. 1986; Ferembach 1976b; Howell 1978). The fragmentary material was associated with an Aterian assemblage and are dated in excess of 20,000 years BP (Brauer 1984a; Clark 1982; Debenath et al. 1986; Ferembach 1976b; Howell 1978). Dar-es-Soltan 5, from the site ofDar-es-Soltan II, is the most complete and only published individual from this sample (Ferembach 1976b). This anterior cranium displays a moderately low sagittal curvature of the frontal bone, although the supraglabellar region is quite steep, a prominent glabella and a well-developed supraorbital torus that appears to thin laterally. The flat face is relatively broad and short, with large rectangular orbits, a wide interorbital area, a low nasal height and some degree of alveolar prognathism (Brauer 1984a; Ferembach 1976b; Howell 1978). The associated mandible displays moderate chin development, a relatively short and broad ascending ramus, large teeth and no retromalar space (Debenath et al. 1986; Ferembacb 1976b).

Temara 2 is an occipito-parietal fragment (Brauer 1984a; Ferembach 1976b). It is quite rounded in profile lacking any bunning, and has a gracile nuchal plane. The Temara 1 mandible was initially assigned to the beginning of the last interglacial (Presoltanian) period (Biberson 1963), but more recent investigations at the site, during which Temara 2 was recovered, resulted in the hominid material being assigned to the Upper Aterian (Roche and Texier 1976), dating to at least 30,000 years BP (see below). However, Briggs (quoting a paper by G. Hubert) stated that The mandible [Temara 1], included in a block of pink sandstone, was found re-deposited in the last level containing Aterian. (1968:380)

Brauer (1984a) suggested that Dar-es-Soltan 5 was associated with the Upper Aterian and dated to around 30,000 years BP, although he has more recently suggested that Dar-es-Soltan II is at least 40,000 years old and possibly 50-70,000 years old (Brauer 1992). Stringer et al. (1984) stated that it is early Aterian, and so dates in excess of 40,000 years BP, and Clark ( 1982) associated it with the lower Aterian which he dated to greater than 30,000 years BP. McBrearty and Brooks (2000) gave Dar-es-Soltan a preferred age estimate of 60-90,000 years BP based on dates for Aterian material elsewhere in north Africa.

Brauer ( 1992) has suggested that the Temara hominids are at least 40,000 years old and possibly 50-70,000 years old. McBrearty and Brooks (2000) cite preferred age estimates for the Mugharet el-'Aliya material of 130-40,000 years BP based on associated artefacts and 60-90,000 years BP based on dates for Aterian material elsewhere in north Africa.

Dar-es-Soltan 5 was not fully published and so could not be analysed. Brauer (1992:Figure 3) included Dar-es-Soltan 5 in a principal components analysis where it was placed near Cro-Magnon 1 and not far removed from other European Upper Palaeolithic crania and late Pleistocene or early Holocene sub-Saharan African material. On a principal components analysis of facial measurements Brauer and Rimbach ( 1990:Figure 1) found Dar-es-Soltan 5 to be well separated from other late Pleistocene north African material, European Upper Palaeolithic crania, Skhul 5, Jebel Qafzeh 6 and Neanderthals. When facial and vault measurements were used, Dar-es-Soltan 5 was found to be near Cro-Magnon 1 and not far removed from sub-Saharan African material (Brauer and Rimbach 1990:Figure 2). Again Dar-es-Soltan 5 was well separated from late Pleistocene north African material from Taforalt and Afalou-bou-Rhumrnel. On this analysis Dar-es-Soltan 5 was also well separated from Skhul 5 and to a lesser extent Jebel Qafzeh 6. On a discriminant function analysis of facial measurements Brauer and Rimbach (1990:Figure 7) found Dar-es-Soltan 5 to be placed within the area of overlap between European Upper Palaeolithic and north African material.

The Temara hominid material displays both archaic and more modem features. The mandible, especially, has archaic features and so its placement in an earlier period, as initially suggested, cannot be totally discounted (Wolpoff 1980a). Tobias suggests that Temara 1 ...should clearly be regarded as a late and somewhat evolved derivative of the "pithecanthropine" mandibles of earlier African times. (1968:184) whereas, Vallois and Roche (1958) saw it as a preNeanderthalian comparable in evolutionary stage to the Steinheim cranium and the Montmaurin mandible from Europe. Brauer (1984a) provisionally placed Temara 1 in his Homo sapiens Grade 2a (Neandertaloid Homo sapiens ), and Temara 2 in his Homo sapiens Grade 3 ([early] anatomically modem Homo sapiens), which emphasises their morphological differences. McBrearty and Brooks (2000) placed Temara, along with Dar-es-Soltan, Omo 1 and the material from Klasies River Mouth and Border Cave into their Group 3. 67

Afalou type.. . which basically represents a Cro-Magnon type", and so represents the type of hominid that moved into north Africa and replaced the local populations .

Dar-es-Soltan 5 has affinities with the Jebel Irhoud crania, although its face appears to be smaller and less prognathic than that of Jebel Irhoud 1 (Brauer 1984a; Ferembach 1976b; Stringer et al. 1984; Wolpoff 1980a). Wolpoff (1980a) suggested that it resembled the male crania from Jebel Qafzeh, Israel. Although robust, Dar-es-Soltan 5 is like most of the other Aterian hominids in that it is modem in overall morphology. Stringer et al. contend that Dar-es-Soltan 5 ...appears to show affinitie s to both the more archaic Irhoud and the "modem" Afalou samples from North Africa . (1984:120) and Smith stated that Dar-es-Soltan 5 ...can be viewed as a good morphological intermediate between archaic H. sapiens and later modem H. sapiens (like the Mechta Afalou sample) in North Africa and as evidence supportin g a morphological continuum between archaic and modem H. sapiens in this region. (1985:205)

Thoma (1973 , 1978a, 1985) suggested that a migration into north Africa from east Africa (possibly an Omo 2 type) is responsible for the origin of modem hominids in the region. It is interesting to note that the above scholars see two morphologically very different hominids as representing the type that migrated into north Africa and replaced the local forms. Why and how this occurred is not elaborated upon. Jaeger (1981) identified two chronologically distinct groups in north Africa , and supported a younger date of 220,000 years BP to 150,000 years BP for evolved Homo erectus fossils such as Sale, Sidi Abderrahman and Rabat, which brings them much closer to the earliest modem skeletal material from the region. He stated that ....a progressive evolution of the group into Homo sapiens is unlikely since it would have had to occur in a very short time. (1981: 159)

McBrearty and Brooks (2000) placed Dar-es-Soltan , along with Temara, Omo 1 and the material from Klasies River Mouth and Border Cave into their Group 3.

However, an earlier dating of the evolved Homo erectus material from north Africa is more probable on current evidence, and this would negate Jaeger's proposition .

Discussion

P .E. Smith contends that ...the early blade-tool industries and the earliest modem men in North Africa owed more to the diffusion from outside of styles, techniques and people than to purely local developments. (1982:404)

There are two major explanations as to the origin of modem humans in north Africa. One, the Replacement Hypothesis, argues that modem humans moved into the region and replaced the local inhabitants, whereas the other, the Multiregional Hypothesis, sees morphological continuity throughout the evolutionary sequence in north Africa. The latter explanation is currently the most widely accepted for this region.

The north African hominid sample has also been seen as supporting the Multiregional Hypothesis, which postulates local in situ evolution of modem humans.

Brauer, however, supported the Replacement Hypothesis, stating that ...a.m. humans spread from the East African area into the North ...The replacement of the North African Neandertaloids also appears to have occurred in the course of this postulated expansion. The more recent human remains from the Upper Aterian (c 30 ky BP) of Morocco may possibly represent evidence of this replacement. (1984a:395) and that The middle to late Upper Pleistocene finds from Northern Africa ....when considered in the light of the fossil finds from Eastern and Southern Africa, indicate that it is highly unlikely that a direct development to modem Homo sapiens took place in this north-western part of Africa . For populations with N eandertaloid characteristics (eg. Mugharet el'Aliya) were still living there some 30,000 years ago, while, much earlier , anatomically modem humans had already spread throughout sub-Saharan Africa . ( 1984b: 157) Brauer (1984b:158) argued that the Omo 1 calvaria from Ethiopia "exhibits strong morphological similarities to the

Briggs concluded ...the hominid remains ...represent links in a North African chain of hominid evolution which developed locally and remained independent of all external influence ... (1968:377) Tobias contends that Several lines of evidence suggest a transition from Middle Pleistocene pithecanthropines to the Upper Pleistocene neandertaloids . Most notable are the mandibular links, for although evolved in character the Upper Pleistocene mandibles ...still betray a complex of traits which are to be found more fully developed in the Middle Pleistocene mandibles . Cranially too, there are pointers tending to close the gap. (1968:189) Jelinek stated I hold that the north-African specimens represent the local evolution leading to Homo sapiens sapiens. (1980b:113)

68

She proposed that 50,000 years ago an Aterian group migrated from north Africa across the Mediterranean Sea to southern Europe, which "would account for the sudden appearance of the Cromagnoids in Europe" (Ferembach 1985:395). The "Aterians" who remained in north Africa are suggested to have evolved at a slower rate than those in Europe, who, at around 24,000 years ago, migrated back to north Africa and gradually replaced the indigenous "Aterians" (Ferembach 1985, 1986). A terminal Pleistocene migration by people from the Levant who assimilated with and/or drove out the local inhabitants is also proposed by Ferembach (1985, 1986). However, there is little support from the archaeological record or the skeletal sample that supports Ferembach's (1985, 1986) argument for transMediterranean movements of the north Africans who produced the Aterian indu~try.

Tobias (1967; 1968) also identified similarities between the Haua Fteah mandibular fragments and mandibles from subSaharan Africa including that from the Cave of Hearths (Tobias 1971), but as discussed earlier, the comparisons are very general and the features have a wide distribution. Wolpoff suggested that It is easy to see continuity with earlier samples. A population with the features of ...Rabat could develop into one with the Jebel lrhoud features through facial reduction (especially in the maxillary area), frontal expansion, and some expansion of the upper portion of the occipital bone. ( 1980a:262) and that the north African Mesolithic samples from sites including Wadi Haifa, Afalou-bou-Rhummel and Taforalt, are robust versions of more recent north African samples.

Lithic Material

Trinkaus argued that although the evidence is weak ...the fossil evidence is consistent nonetheless with a model of gradual change from the Acheulian associated Rabat, Sidi Abderrahman and Temara specimens to the Mousterian associated Irhoud remains. (1982a:307)

If anatomically modem humans moved into north Africa there should be evidence of a break or change in the archaeological record.

The Aterian As alluded to above, there are problems with the chronological placement of the Aterian. There is said to be evidence to suggest that the Aterian was a direct derivative of, contemporary with, and the successor of the Maghreb Mousterian (Clark 1982). Ferring, however, contends that It [the Aterian] had its origin in the Mousterian of North Africa, sometime in the early Wiinn, with radiometric evidence for its spread from its place of origin in excess of 40,000 years ago. (1975: 121) Clark outlined the dates obtained for the Aterian throughout north Africa and stated that ...the Aterian ...belongs to a time from more than 45,000 years BP to an absolute minimum of 30,000 years ago and more likely all the Aterian is more than 40,000 years ago ... (1982:3) Clark ( 1982) dated the Aterian to between ?70,000 years BP and ?35,000 years BP. Debenath et al. (1986) dated the Aterian in excess of 20,000 years BP on the bases of an Upper Palaeolithic layer at Taforalt that overlies Aterian levels which is dated to 21,900+/-400 years BP (Gif-2585), and because at Temara, where there is evidence of a hiatus of at least 5,000 years between Upper Palaeolithic and Aterian levels, the former is dated to 14,469+/-200 years BP (Laboratory number not given). Debenath et al. ( 1986) also recorded that at Temara the Middle Aterian had produced dates of 23,700+/-1000 years BP (Gif-2585) and 24,500+/600 years P (Gif-2582). Taforalt has also produced dates for the Aterian of greater than 40,000 years BP (Clark 1982; Debenath et al. 1986). The Moroccan site of Ain Maarouf has provided a date of 32,000+/-600 years BP (GrN-3165) for a "typical" Aterian assemblage (Ferring 1975). McBrearty and Brooks (2000:489) suggested that the radiocarbon dates used to place the end of the Aterian at only 20,000 years BP should "probably be treated as infinite". The site of Bir Tarfawi in the western desert of Egypt has dates for the Aterian of > 100,000 years BP produced by the

Stringer et al. noted that From North Africa there is evidence of hominids displaying some signs of local continuity from the early Middle Pleistocene (Ternifine) to the Upper Pleistocene (Djebel Irhoud and Dar-es-Soltan). (1984: 120) Klein and Scott contend that the north African Upper Pleistocene sample ...appears to bridge the morphological gap between archaic and fully modem Homo sapiens better than any other known fossil sample...Together with sub-Saharan human fossils ...the North African ones imply that the roots of fully modem people may lie in Africa. (1986:521) It has also been argued that the north Africa hominid sample documents regional continuity and the in situ evolution of Homo erectus into Homo sapiens, and that it was not until the late Upper Pleistocene that there were migrations into and out of the region. Coon (1962) argued that the ancestors of the modem southern African Khoisan evolved in north Africa until they were forced out of the region by Caucasoids moving in from the east towards the end of the Pleistocene (some mixing between the two groups is also suggested). Coon ( 1962) also postulated a more recent migration from the Levant. Ferembach stated that Man, present in that area [north Africa] at least as early as Homo erectus ...evolves towards the Mousterian Proto-sapiens forms of Djebel Irhoud ...Aterians, archaic Crornagnoids, developed from these last ones. (1985:395) 69

anatomically modem , although robust , humans such as Dares-Soltan 5 had already appeared before the end of the Aterian .

uranium-series disequilibrium and thermoluminescence methods (Clark 1989). McBrearty and Brooks (2000) gave an age range of 60-90,000 years BP for Aterian material in Libya and suggested that the Aterian may be as old as 160,000 years BP at the Dakleh oasis in Egypt.

There may have been some terminal Pleistocene or early Holocene migrations reflected in the variability of the Mesolithic crania from the region (Anderson 1968; Brauer 1978; Briggs 1950, 1955, 1968; Greene and Armelagos 1972), although others suggest continuity throughout this period (Lubell et al. 1984).

P.E. Smith (1982:378) suggested that the Aterian was probably still in existence 30,000 years ago, and possibly could have lasted until around 20,000 years BP, but suggested that "How long the Aterian survived is not known".

There are no Neanderthal remains recovered from north Africa ( contra for example, Brauer 1984a; Ennouchi 1962, 1963; Howell 1978 and Tobias 1968) . El-'Aliya does lack a canine fossa, which is a typical Neanderthal characteristic (Table 5:6), but this appea~s to be a primitive retention as the Rabat maxilla also lacks a canine fossa. During the multivariate analyses Jebel Irhoud 1 did, on occasions, group with the Neanderthals , but as explained earlier, there is only a superficia l resemblance between Jebel Irhoud 1 and the Neanderthals in the posterio r vault , while the face of Jebel Irhoud 1 does not display any characteristic Neanderthal features (Table 5:6).

The Aterian, therefore , appears to date from greater than 40,000 years BP down until possibly 20,000 years BP, with most occurrence s older than at least 30,000 years BP (Camps 1975; Clark 1982, 1989; Debenath et al. 1986; Ferring 1975; Smith, P .E. 1982).

The Dabban P.E. Smith's (1982) suggestion of an influx into the region from outside is based , predominantly , on the contention that the Dabban industry from the Jebel el-Akkdar, which replaced the local Levalloiso-Mousterian industries sometime after 40,000 years BP, did not have local origins. Smith stated that the Dabban industry ...may represent a true diffusion from outside the Jebel el-Akkdar, perhaps from south-west Asia ...and possibly ...by men of modem sapiens type. (1982:353) The Dabban industry is only found in a restricted part of north Africa , with P.E. Smith (1982:354) stating that "It is curious that there is no certain case of Dabban occupation known in any other part of North Africa" . The Dabban industry cannot be used to demonstrate a major influx of people into north Africa around 40,000 years BP because if it marked such an influx there should have been a much wider distribution of it throughout the region and not a restricted occurrence in the Jebel el-Akkdar . Also , to argue that the Dabban industry came from outside of north Africa there must be antecedents for it elsewhere , and evidence of its existence outside of this region. This evidence has not been forthcoming. All that P.E . Smith (1982) has suggested is that "perhaps" the Dabban industry came from southwest Asia and that it was "approximately contemporary" with Upper Palaeolithic blade industries from Europe and southwest Asia.

No detailed list of "clade features" that document "regional continuity" in north Africa have been proposed . General similarities in mandibular size and robustness, and dental form such as wrinkled enamel on molars, slight taurodontism , and slightly incisiformed canines, have been suggested ( cf. Brauer 1984a; Briggs 1968; Coon 1962; Hublin and Tillier 1981; Wolpoff 1980a). Stringer et al. (1984) have suggested some characters of the splanchnocranium shared by Jebel Irhoud 1 and Dar-esSoltan 5, including a low nasal height in relation to the upper facial breadth , suggest evolutionary continuity. One could add a lack of mid-facial prognathism , which links these crania to mandibular remains from the region such as the Haua Fteah material. It is also possible to document a continuous single-directional pattern of cranial and dental change reflecting a decrease in overall cranial robustness (Carlson and van Gerven 1977; Chamla 1981; Wolpoff 1980a). As Smith observed for north Africa ...there is no convincing evidence, either in the hominid fossil or cultural records, of significant intrusive elements during the period of transition to modem H. sapiens . (1985:205) North Africa would seem to demonstrate the in situ evolution of anatomically modem humans . Due to the fragmentary nature of much of the earlier hominid remains, the lack of chronological controls on many of the finds, and a scarcity of relatively complete archaic Homo sapiens crania, the evolutionary sequence is partially obscured. In many instances "grade" similarities between regions dominate any "clade" similarities. The latter (if they actually exist, cf. Tattersall 1986), remain to be clearly identified for north Africa , but this does not negate the evidence of morphological and cultural continuity and the lack of evidence of major external influence. This does not however, discount the possibility of gene flow from outside of north Africa .

Lubell et al. (1984) have identified similarities in the EpiPalaeolithic industries throughout north Africa, which is more compatible with an in situ evolutionary development.

Conclusion The sample of hominid remains from north Africa can be interpreted as documenting an evolutionary progression from Homo erectus through to anatomically modem Homo sapiens without significant influx of people from other areas. There is a major hiatus after the Aterian occupation at many sites, from about 30,000 years BP to 15,000 years BP, but this may be climatically induced - an arid phase caused by the onset of the last glacial maximum (Ferring 1975). In any case, 70

CHAPTER FOUR. WESTERN ASIA Introduction

Dmanisi

For this Chapter, western Asia encompasses three major regions, the Levant (Shanidar Cave from Iraq has been included in this region), central Asia, and south Asia (Figs. 4:1, 4:2, 4:9, 4:12, Table 4:1).

The site of Dmanisi, Georgia (Fig. 4.1) has produced four well-preserved crania, mandibular, postcranial material and dental remains, Oldawan Mode 1-type stone tools and faunal material (Gabunia and Vekua 1995; Gabunia et al. 2000; Gibbons 2003a; Vekua et al. 2002). Hominid occupation at the site is correlated to a period just following the Olduvai-Matuyama polarity boundary and is given an age of approximately 1.7 million years ago. The underlying Masavera Basalt has provided an age of 1.85 million years ago (Gabunia et al. 2000). The faunal remains from the site support a latest Pliocene-earliest Pleistocene age (Gabunia et al. 2000; Vekua et al. 2002). Whilst there is morphological variation between the crania, they can be attributed to Homo erectus, although the mandible has been classified as Homo georgicus (Gabunia and Vekua 1995; Gabunia et al. 2000; Gibbons 2003a; Vekua et al. 2002). The smallest of the three crania, D2700, is said to also show similarities with African Homo habilis material (Vekua et al. 2002), but parallels can also be found with the Homo erectus calvaria from Ileret, Kenya (Gibbons 2003a). In general, the crania are relatively small with low vaults, thin but prominent brow ridges, marked postorbital constriction, maximum breadth

Westem Asia has been an area of major interest to studies of human evolution because it has produced relatively good samples of archaic and modem hominids. The Mount Carmel hominids from Israel have been debated the most of all of the hominid remains from western Asia, with suggestions that they demonstrate in situ evolution from Neanderthals to anatomically modem humans, two separate evolutionary lines with one being Neanderthal and the other modem humans, or hybridisation between Neanderthals and anatomically modem humans. The focus on the western Asian region has intensified over the last few decades with revised dating of key sites and new discoveries. Howell ( 1998) details some of the earlier archaeological sites from the region that will not be discussed here.

Table 4:1. Western Asian hominids examined, abbreviations for Figures and Tables, and source of data. Hominid Amud 1 Amud2 Amud7 Dederiyeh Dmanisi material Hotu2 Jebel Qafzeh 6 Jebel Qafzeh 9 Jebel Qafzeh 11 Jebel Qafzeh Aurignacian material Kebara 1 Kebara 2 Ksar 'Akil 1 Mezmaiskaya Nahal 'En-Gev 1 Narmada Shanidar 1 Shanidar 2 Shanidar 4 Shanidar 5 Skhiil 1 Skhiil 4 Skhiil 5 Skhiil 9 Starosel'e Tabiin Cl Tabiin C2 Teshik-Tash Zuttiyeh

Original Or Cast Original Published Published Published Published Cast Original Original Published Cast Published Cast Cast Published Published Cast Cast Published Published Cast Original Original Original Original Published Original Original Cast Original

71

Abbreviations Al --

---

-H2 JQ6 JQ9

---

-----

-Na Shl Sh2 Sh4 Sh5

-Sk4 Sk5 Sk9

-TCl TC2 TT

--

Principal Source Of Data Original Suzuki 1970b Rak et al. 1994 Dodo et al. 1998 Gabunia et al. 2000; Vekua et al. 2002 Cast Original Original Tillier 1984 Cast Smith and Arensburg 1978 Cast Cast Golovanova et al. 1999 Arensburg 1977 de Lumley, M-A and Sonakia 1985 Cast, Trinkaus 1983a Trinkaus 1983a Trinkaus 1983a Cast, Trinkaus 1983a Original Original Original Original Alexeyev 1976 Original Original Cast Original

MEDITERRANEAN

SEA

Amud Zuttiyeh• • L. Tiberias

'Ubeidiya • Oafzeh

•Jebel

Figure 4:1. Map showing the Levantine sites mentioned in the text.

sites discussed in this section are located in Israel, the southern Levant, although fragmentary hominid remains from the relevant period have been recovered from throughout the Levant (Fig . 4: 1). The hominid material from Shanidar Cave in Iraq has, for convenience, also been dealt with in this section (Fig. 4:9).

near the mastoid region and where preserved (on D2282 and D2700) the facial regions are relatively small. The Dmanisi material is significant because it demonstrates the early migration out of Africa of small-brained hominids. What is still to be answered is whether this material represents the ancestor of Asian Homo erectus material and later European hominids.

Galilee-Mugharet el-Zuttiyeh

The Levant

A hominid frontofacial fragment was recovered from Zuttiyeh (Robbers') cave in the Wadi el'Amud , upstream from the northwest end of the Sea of Galilee, Israel (Plate 16; Fig. 4:1; Keith 1931; Oakley et al. 1975). The fragment consists of a robust frontal squama with the right malar and sphenoid bones (Hrdlicka 1930; Keith 1931; Trinkaus 1984a; Vandermeersch 1981c, 1982).

The site of 'Ubeidiya , Israel (faunal remains, lithic material and a hominid molar, and two lower incisors), demonstrates the presence of hominids in the Levant in excess of 1 million years ago (Belmaker et al. 2002) . However , relatively complete hominid remains have only been recovered from more recent sites. The majority of the

72

Age in kyrs B.P.

p

40

STAROSEL'E ?t

f;J AMUO 1

~

50

TESHIK-TASH? t

~~ SHANIDAR

KEBARA

SHANIDAR 2 & 4 ? t+

100

~

JEBEL OAF

BUN ?t ♦

150

SKHUL

Q

?t

ZUTTIYEH ?+

Figure 4:2. The probable chronological position of the Western Asian hominid sample . 73

the breadth of the upper face in general, and the interorbital region in particular . A similar grouping with Kabwe 1 was found in earlier analyses (Habgood 1982, 1984b; Habgood and Walker 1986).

The moderately thick frontal bone displays a relatively steep supraglabellar portion and then gently recedes. The frontal is relatively narrow, which suggests a high degree of postorbital constriction. The supraorbital torus, which is separated from the frontal by a marked ophryonic groove, is well-developed. Vandermeersch ( 1989) described the brows as being divided into superci/iaris and supraorbitalis segments. Whilst there is some medial flattening on the right side of the supraorbital torus, this is most likely due to a pathological lesion on the superior surface of the torus (Simmons et al. 1991). The left supraorbital is not separated into medial and lateral segments. The supraorbital torus does not really taper laterally. The glabella does not protrude to the same extent as the supraorbital torus and the nasion is not recessed. The right malar is delicately constructed when compared to the supraorbital region. The high nasiofrontal angle of Zuttiyeh indicates that the upper portions of the nasal bones, at least, and probably the lower mid-facial regions, did not project markedly (Stringer 1978, 1989; Trinkaus 1983a; but see Simmons et al. 1991; Smith and Paquette 1989). It also indicates that the upper face was relatively flat. The interorbital region is wide. The quadrangular right orbit is not overly large. The orbital margin is quite thick and the upper face appears to have been relatively broad .

This grouping is compatible with the overall morphological configuration of Zuttiyeh , which does display similarities with these two crania. All three display well-developed supraorbital regions, with Zuttiyeh and Kabwe 1 having tori that do not taper laterally. Their upper faces are broad and robust, especially the interorbital region. The mid-facial region of Zuttiyeh is relatively flat like Skhiil 5, but different from the mid-facially prognathic face of Kabwe 1 (Stringer 1978). Both Kabwe 1 and Skhul 5 are more prognathic than Zuttiyeh appears to have been (Stringer 1978, 1989; Trinkaus 1983a; but see Simmons et al. 1991). The frontal bone of Zuttiyeh is higher than that of Kabwe 1, but not as high as Skhiil 5. In this feature Zuttiyeh is closer to Laetoli H18 , Florisbad, Jebel Irhoud 1, Amud 1 and Skhiil 4 and 9. Morphologically, Zuttiyeh could be placed between Kabwe 1 and Skhfil 5, but in general, it displays more overall resemblance to the former in that it retains more archaic features. Vandermeersch (1981c, 1982, 1989) suggested that Zuttiyeh lacked any specialised Neanderthal features {Table 5:6) and that the differences between Zuttiyeh and the Neanderthals from western Asia are greater than those with the material from Skhul and Qafzeh. He concluded that Zuttiyeh was the most ancient example of Homo sapiens sapiens in the region. Stringer (1978, 1989) has also identified a distinct difference between Zuttiyeh and the "classic" Neanderthals.

The Zuttiyeh hominid was derived from an AcheuloYabrudian facies context (A.J. Jelinek [1982a] included this facies in his Mugharan Tradition) at the site (Gisis and BarYosef 1974). The Acheulo-Yabrudian assemblage, which is similar to the lithic material from Tabiin Layer E, is restricted to the northern and central Levant (Bar-Yosef 1989). The Acheulo-Yabrudian, was placed between 140,000 and 90,000 years BP (Jelinek, A.J. 1982a, but see Mercier et al. 1995), however, with refined dating and lithological analyses it has been placed within oxygen isotope stage 6 dating to either 130-190,000 years BP (Table 5:1; Bar-Yosef 1989), or even earlier to 250-350,000 years BP (Bar-Yosef 1998; Mercier et al. 1995) or 200-380,000 years BP (Barkai et al. 2003). Travertine deposits underlying the AcheuloYabrudian level at Zuttiyeh have provided uranium series (Th/U) dates of 148,000+/-6,000 years BP and 164,000+/21,000 years BP, while the layers above the AcheuloYabrudian at the site have provided dates of 157,000+/13,000, 106,000+/-7,000, 97,000+/-13,000 and 95,000+/10,000 years BP (Bar-Y osef 1989, 1998:Table 1; Schwarcz 1980, 1982; Valladas et al. 1998). Whilst there is some stratigraphical inconsistency , these dates support the younger dating scheme for the Acheulo-Yabrudian. Thermoluminescence dating of burnt flints from Tabiin, if correct, would place the transition from Acheulo- Yabrudian to Mousterian industries at between 250-270 ,000 years BP (Mercier et al. 1995), but other radiometric dating at the site suggests an indicative age of 150-250,000 years BP (Griin et al. 1991; McDermott et al. 1993). A date of around 150,000 years BP would be compatible with the available data, although an earlier date in excess of 200,000 years BP is still possible (Fig. 4:2).

Hills and Brothwell ( 1974) found Zuttiyeh to fall near the Neanderthals and suggested that their results supported the view that Zuttiyeh was a "classic" Neanderthal. Similar conclusions have been reached by other investigations. Smith et al. (1989) contend that Zuttiyeh is similar to European Neanderthals in frontal morphology and "other details", although it lacks midfacial (midsagittal) prognathism. Trinkaus ( 1983a:465) argued that Zuttiyeh was "probably best considered as a late early Neanderthal", and that it exhibits traits which are symplesiomorphies for Upper Pleistocene hominids in general (Trinkaus 1988; see also Smith et al. 1989). Trinkaus (1982a, 1983a) also suggested that Zuttiyeh displayed morphological similarities to the fragmentary Shanidar 2 and 4 individuals. Shanidar 2 and Zuttiyeh preserve mostly different regions of the cranium: Shanidar 2 the posterior cranium and lower face and Zuttiyeh the upper anterior cranium. Therefore, comparisons between the two individuals are difficult, however there do not seem to be any marked similarities. Shanidar 4 displays more similarity to Zuttiyeh in that it preserves a portion of a robust supraorbital torus that appears not to have tapered laterally, and it has a steep supraglabellar region, although the remainder of the frontal is much flatter than that of Zuttiyeh (Trinkaus 1983a). Simmons et al. (1991) did not find a close link between Zuttiyeh and Shanidar 1 and 5.

Due to the fragmentary nature of Zuttiyeh only a limited number of measurements were available , with a number of these being estimates . Zuttiyeh was found to group with Kabwe 1 and Skhiil 5. There were high loadings on the variables that document the form of the frontal squama and

Simmons et al. ( 1991) undertook a multivariate analysis of the frontal so as to compare Zuttiyeh with other hominids 74

archaic Homo sapiens). They also contend that, due to the lack of these east Asian clade features, it is unlikely that Middle Pleistocene · hominids from Europe or Africa were ancestral to Zuttiyeh.

from western Asia. They found that the Zuttiyeh frontal grouped with Amud 1 on the plots from the principal components analyses of log size and shape and log shape and the cluster analysis (Simmons et al. 1991 :Figures 1, 2 and 4 ). It is worth noting that Zuttiyeh was also not far removed from the Jebel Qafzeh 6 and Skhiil 4 and 9 hominids. Simmons et al. ( 1991) concluded that Zuttiyeh was a logical ancestor for western Asian Neanderthals and not the hominids from Skhiil and Jebel Qafzeh. In a pairwise clustering analysis based size and shape distance statistics Corruccini (1992:Fig. 2) also found Zuttiyeh to group with Amud 1 and then cluster with western Asian and European Neanderthals. Suzuki ( 1970b) did not identify close morphological similarities between Zuttiyeh and Amud 1. For example, Suzuki ( 1970b) identified that the supraorbital torus of Zuttiyeh was heavier than that of Amud 1, whereas the malar was more gracile. Suzuki (1970b:189) concluded, "As a whole, the Galilee man is a more archaic type than Amud".

Thoma (1964, 1973) also argued for a link between western and eastern Asia, but was of the view that western Asian Neanderthals (Amud, Shanidar and Teshik-Tash) formed part of the ancestry of eastern Asian Mongoloids (see also Pichardo 1978), whereas Sohn and Wolpoffs model is based on gene flow ( or population movement?) from east Asia into western Asia. There are also similarities in the shape of the orbits and supraorbital region between Zutteiyeh and the Narmada calvaria. Holt and Waddle ( 1991) undertook a multivariate comparison of Zuttiyeh with Homo erectus material from Africa and east Asia and archaic Homo sapiens material from Europe and Africa. They found that metrical comparisons did not support an east Asian origin for Zuttiyeh as opposed to an African origin. The morphological similarities between Zuttiyeh and African Middle Pleistocene material are greater than proposed by Sohn and Wolpoff (1993), but there are also morphological differences (Chapter 2 and 7; Habgood 1992).

Zuttiyeh displays some morphological similarities to Tabun Cl in that they both have a prominent ophryonic groove behind a supraorbital torus that does not taper laterally and only moderate facial prognathism. Simmons et al. (1991:Figures 1, 2 and 4) found Tabiin Cl to be not far removed from Zuttiyeh. Sohn and Wolpoff (1993) also identified morphological similarities, especially around the region of the nasal root, between Zuttiyeh and Tabun Cl. However, they found few similarities with Amud 1, the only other Neanderthal in the comparisons, and note numerous similarities with the material from Skhul and Jebel Qafzeh, especially Jebel Qafzeh 6 (Sohn and Wolpoff 1993:Table 5).

Whilst it is difficult to conclusively establish the evolutionary status of Zuttiyeh, the present investigation indicated a distinction between Zuttiyeh and the Neanderthals from both Europe and western Asia. Zuttiyeh does not appear to be a Neanderthal, but it is clearly an archaic form and not an early representative of Homo sapiens sapiens. Rak (1986) has proposed that Zuttiyeh, and other north African and Levantine material, has what he proposed was a generalised facial form.

In some morphological features Zuttiyeh also recalls the calotte from Maha, China (Plate 16). These include the curvature of the frontal squama, the degree of postorbital constriction and the flattish upper face, although Zuttiyeh has larger non-tapering brow-ridges, a more prominent ophryonic groove and a more rectangular orbit. W olpoff ( 1989) also identified parallels with the Homo erectus remains from Zhoukoudian. Sohn and Wolpoff (1993) undertook a morphological comparison of this material with Zuttiyeh. Univariate metric comparisons did not separated the Zuttiyeh frontal from the other Levantine hominid material, but Sohn and Wolpoff (1993) argued for strong morphological affinities with the material from Zhoukoudian when metrical features are combined on bivariate plots. For example, maximum frontal breadth and frontal length, frontal curvature index and nasion projection, and projection and thickness of the supraorbital torus. When combined these metrical comparisons are used to argue for similarities between Zuttiyeh and the Zhoukoudian material in overall frontal and supraorbital torus size and shape (Sohn and Wolpoff 1993). Whilst acknowledging that the Zuttiyeh frontal is not a "west Asian version of the Zhoukoudian folk", and that there are important differences such as thinner squama and lack of frontal keeling on Zuttiyeh, Sohn and Wolpoff (1993:340) suggested that the morphological similarities are "striking". These "striking similarities" include the form of the supratoral sulcus, rounded supraorbital contour, the flattened nasal region and midfacial flatness. Sohn and Wolpoff (1993) concluded that Zuttiyeh preserved a number of clade features from east Asia that are also present in varying combinations in latter Levantine hominids (both Levantine Neanderthals and

Stringer et al. ( 1979) placed Zuttiyeh in their Homo sapiens Grade 2 or 3.

Mugbaret es-Skbiil At least 10 and up to 26 individuals were recovered from Layer B, Levalloiso-Mousterian levels, at the cave of esSkhul in the Wadi el-Mughara, Mount Carmel, Israel (Fig. 4:1; Garrod and Bate 1937; Hovers et al. 1995; Oakley et al. 1975). Layer B was subdivided into an upper B 1 unit and a lower B2 unit (Garrod and Bate 1937). Possibly seven of the Skhiil individuals appear to be intentional burials that were clustered on the terrace in front of the cave (Belfer-Cohen and Hovers 1992; Garrod and Bate 1937; Hovers et al. 1995:Fig. 1; Table 2; Ronen 1976:Fig 1). Only the three most complete crania, Skhul 4, 5 and 9, will be dealt with here, but descriptions of the other material may be found in McCown and Keith ( 1939). The postcranial material from Skhul is robust but essentially modem in configuration and displays similarities with the Jebel Qafzeh sample and contrasts with western Asian Neanderthals (McCown and Keith 1939; Trinkaus 1983a, 1983b, 1993, 1995; Trinkaus et al. 1998). The Skhiil individuals can be typified as possessing linear bodies with narrow trunks and long robust limbs (Trinkaus 1995 and references therein). 75

Skhul 1 is the skeleton of an infant aged approximately 4 years old (Oakley et al. 1975). Skhul 1 has squared-off superior orbital margins , clearly distinguishable medial and lateral supraorbital segments , developed parietal eminences , no lambdoidal flattening on parietals , rounded foramen magnum, well-developed mastoids and absence of juxta mastoid eminences , which align it with archaic Homo sapi ens and not Neanderthals (Minugh-Purvis 1998; Rak et al. 1994; Tillier 1998). Tillier (1998) did note that the mandible of Skhul 1 displayed archaic features including enlargement of the dental arch and anterior alveolar flattening , but these features were also found on immature specimens from Jebel Qafzeh and the Upper Palaeolithic associated child from Ksar 'Akil in Lebanon . Schwartz and Tattersall (2000) have described the symphyseal region of the Skhfil 1 mandible as featureless .

2000 and discussion above) , a slight retromolar gap, mental foramen positioned under the fourth premolar and does not have , like Skhiil 4 and Jebel Qafzeh 9, the horizontal-oval mandibular foramen pattern . Skhiil 9 consists of a fragmentary adult male skeleton (McCown and Keith 1939). The cranium is quite long and low with little postorbital constriction . The supraglabellar region of the frontal bone is quite steep, while the parietals display postbregmatic and pre-lambdoidal flattening. The occipital is rounded and has an occipital torus ( contra Santa Luca [1978], cf. note above), but no suprainiac fossa. The supraorbital region is well-developed with the supraorbital trigone distinguishable from the superciliary arch. The dating of the Skhiil hominid material has caused much debate since they were first discovered . The hominid material was initially assumed to be of the same age as the hominids from Tabiin Layer C because of the nature of the Levalloiso-Mousterian artefact s and the associated fauna (Garrod and Bate 1937; McCown and Keith 1939). Higgs ( 1961; Higgs and Brothwell 1961), studying faunal remains from the Mediterranean region , suggested that Skhiil was possibly 10,000 years younger than Tabiin Layer C, a scenario that had also been proposed by Howell (1958) . A reinterpretation of the Tabiin fauna! sequence indicated that it was not especially useful in elucidating the chronological relationship of the site with Skhiil (Jelinek et al. 1973). A comparison of the lithic material from Skhiil with that from Tabiin was interpreted by Jelinek (Jelinek , A.J 1982a, 1982b) as indicating that the Levalloiso-Mousterian lithic material from Skhiil Layer B was late within the Middle Palaeolithic sequence in the Wadi Mughara and that the material from Skhiil Layer B2 may be contemporary with the "chimney" deposits at Tabiin, while the Skhiil Layer B 1 material postdated these deposits . There is considerable heterogeneity within the lithic assemblage of Layer B at Skhiil, and there are typological comparisons with Tabiin B-type and C-type (Bar-Yosef 1992, 1998; Garrod and Bate 1937; Jelinek, A.J 1982a, 1982b).

Skhul 4 is an adult male skeleton (McCown and Keith 1939). It has a long and relatively low cranium with little postorbital constriction . The frontal however, is quite steep, especially the supraglabellar region. There is some postbregmatic and pre-lambdoidal flattening , but no marked occipital angulation or protrusion . The morphology of the occipitomastoid region is non-Neanderthal in overall configuration in that there is no occipital torus or suprainiac fossa and the mastoid process is large. The face is quite large and displays marked alveolar prognathism . The orbits are rectangular in shape, the nasal aperture is large, and the nasal bones project. The malars are not inflated, but do not display a distinct canine fossa. The supraorbital region is prominent with the superciliary arch distinguishable from the supraorbital trigone. The mandible of Skhul 4 has a short, broad ramus with a slight retromolar gap and the mental foramen located under the first molar. The Skhfil 4 mandible, along with Skhfil 5 and Jebel Qafzeh 9, does not have the horizontal -oval mandibular foramen form . The symphyseal region displays a distinct protrusion that has been classified here as a well-developed mental eminence or chin. Schwartz and Tattersall (2000:380) described the symphyseal region on Skhul 4 and 5 as having "a subalveolar depression from which emanates a low teardrop -shaped bulge that flows into the surrounding bone" , as opposed to the inverted "T" delineated by mental fossae which they found typified modem Homo sapiens.

Revised aspartic acid racemisation dates have been obtained from four of the Skhiil hominids (Masters 1982). These dates are 39,000 and 40,000 years BP from Skhiil 5, 52,000 and 54,000 years BP from Skhul 6, 45,000 and 46,000 years BP from Skhiil 7, and 53,000 and 55,000 years BP for Skhiil 9. There are also problems with this dating technique (Hare 1980) and its use at Skhiil. There were no radiocarbon dates from Skhiil for calibration . The racemisation rate constant obtained from inside Tabiin Cave was used at Skhiil even though the Skhiil material came from an exposed terrace, which could have had a higher average ground temperature and so a higher racemisation rate which , if used, would have produced younger date s. Also , the hominid material from Skhiil had been treated with preservatives that can influence the dates obtained (Masters 1982).

Skhfil 5, an adult male skeleton , is the most complete and widely studied and referred to of the Skhfil hominids (Plates 4 and 6) McCown and Keith 1939; Snow 1953). It has a robust and high vaulted crania with little postorbital constriction. The frontal is high and steep, the parietals are well curved with moderate bossing and the temporals are high and short. There is some pre-lambdoidal flattening and the occipital is generally rounded. There is an occipital torus, without a clearly defined upper margin , that fades laterally, and a shallow suprainiac fossa (contra Santa Luca [ 1978] who used a very restricted definition of these features based on those found on "classic" Neanderthals). The mastoid processes are large and non-tapering . The prominent supraorbital arches, which are separated from the frontal by a marked ophryonic groov e, can be divided into medial and lateral segments . The glabella is also prominent , while the nasal root is depressed . The mid-facial region has been heavily reconstructed (Snow 1953). The orbits are rectangular, and the flat face displays a high degree of alveolar prognathism. The mandible of Skhiil 5 (Plate 8) has a develo ped mental eminence (see Schwartz and Tattersall

Much older dates for Layer B have been obtained by the more recent use of radiometric methods including electron spin resonance , thermoluminescence and uranium series. Electron spin resonance (ESR) dates have been obtained from two bovid teeth from Layer B (Stringer et al. 1989). The exact position of the two bovid teeth within the 2m thick depo sit is not known. The early uptake (EU) ages averaged 76

chronologically separated groupings within Layer B (Garrod and Bate 1937; Ronen 1976:Fig. 1) - Bl[late] Skhul 1, 2, 4, 5; B2[early] Skhul 3, 6-10. Ronen (1976) also suggested that there were two phases of inhumation at Skhiil that were separated by an unknown period of time. The aspartic acid racemisation dating (Masters 1982) can be used to support a separation of inhumations into possibly three phases: Phase 1 at c. 53,000 years BP with Skhul 6 and 9; Phase 2 at c. 45,000 years BP with Skhul 7; and Phase 3 at c. 40,000 years BP with Skhiil 5.

81,000+/-15,000 years BP while the linear uptake (LU) ages averaged 101,000+/-12,000 years BP. Results for sample 521 were: EU - 88,100+/-17,900, 86,100+/-13,100, 94,900+/15,600, 101,00+/-19,000 years BP; and LU - 102,00+/-22,700, 102,00+/-18,100, 109,00+/20.500, 119,00+/-25,100 years BP. Results for sample 522 were: EU - 68,00+/-5,400, 73,000+/-7,000 54,600+/10,300 years BP; and LU - 98,300+/-10,600, 99,900+/-12,400, 77,200+/15,700 years BP.

Whilst these dates are probably too young, the relative ordering of the burials would still be applicable. Bar-Yosef (1992:264), possibly following earlier suggestions (Garrod and Bate 1937; Ronen 1976), postulated that there may have been two burial periods at Skhfil that "represent multiple and sparsely spaced events" - an early period dated at 120,00090/80,000 years BP and a late period dated between 55,00045/43,000 years BP. A sequential burial sequence could reflect a repetitive tradition at Skhiil of inhumation on the terrace in front of the cave (Hovers et al. 1995).

The ages obtained from sample 522 are consistently younger than those from sample 521 and so the bovid tooth used may have been from a stratigraphically higher position within LayerB. McDermott et al. (1993) used the mass-spectrometric uranium-series (U-series) method to provide ages for the two bovid tooth samples analysed by (Stringer et al. 1989). Sample 521DE provided an age of 80,300+/-600 years BP, which is closer to the ESR EU age than the ESR LU result (McDermott et al. 1993). As with the ESR results, sample 522EN provided a younger age than sample 521 - this time 40,400+/-200 years BP, which is significantly younger than the ESR EU and LU ages (McDermott et al. 1993). Two additional samples from Layer B also provided U-series dates in the range 41,400+/-400 to 45,500+/-700 years BP (McDermott et al. 1993).

Attempts are underway to date material directly associated with Skhiil 5 and 9 (Stringer 1998a). When all of the data currently available is considered, an early late Pleistocene age centred around 100,000 years BP for the Layer B at Skhiil seems probable (option 3 above), with the hominids varying in age depending on placement within the stratigraphy (Fig. 4.2). However, as Ronen (1976:31) observed and as is also suggested by the radiometric dating at the site, "the Skhiil sequence stretches over a longer time than originally maintained".

McDermott et al. (1993) concluded that the ESR and Useries dates suggested two faunal ages within Layer B at Skhul. The youngest of a series of calcite layers uranium series date of 79,000+/-4,000 years Stringer et al. 1989), which is consistent uptake ESR averaged age of81,000+/-15,000

Skhiil 4 was found to cluster with the European Neanderthals (Figs. 4:3, 4:4, 4:5, 4:6, Table 4:2), with high loadings on the variables that describe the sagittal contour of the cranium and the breadth of the face (Fig. 4:4, Table A:8). On the analysis of mandibular non-metric traits (Fig. 5: 16) the Skhfil 4 mandible fell near the mandibles of anatomically modem Upper Palaeolithic crania from Europe and Skhiil 5 and was well separated from the Neanderthals from western Asia and Europe and the mandibles from Border Cave and Klasies River Mouth, South Africa. On the ordered similarity matrix {Table 4:7) Skhiil 4 was found to be very similar to Tabiin Cl, similar to Jebel Irhoud 1 and Amud 1 and not very similar to Skhul 5 and Jebel Qafzeh 6 and 9.

has provided a BP (reported in with the early years BP.

Six burnt flints from Layer B at Skhul have provided a thermoluminescence (TL) average age of 119,000+/-18,000 years BP (Mercier et al. 1993; Valladas et al. 1998). This date is in broad agreement with the LU ESR dates obtained from bovid teeth, especially sample 521, but is significantly older than the U-series dates and the EU ESR averaged age. The radiometric dating of Layer B at Skhfil has provided three possible scenarios: 1. An early date of 100-120,000 years BP based ·on LU ESR and TL results; 2. A later date of around 40-80,000 years BP based on EU ESR and U-series results; or 3. An age range of at least 120-80,000 years BP based on all of the methods used.

Because of its completeness, Skhfil 5 was one of the core group of fossils that made up the basic data set that was used Table 4:2. K-means cluster analysis with Skhiil 4, Skhiil 9, Jebel Qafzeh 6, Jebel Qafzeh 9, and Jebel Irhoud 1. Analysis includes 79 .9% of the total within-group variance. Group 1 Skhiil 5, Jebel Qafzeh 9, Combe Capelle, Chancelade, Oberkassel 2, Dolni Vestonice 3.

As Layer B is 2m thick and the provenance within Layer B of some of the material dated is not secure, option three would seem the most appropriate dating scheme to adopt at this time. What also needs to be considered is that at least seven of the Skhfil hominids were probably burials and the material was recovered from different levels within Layer B, with Skhiil 9 for example being recovered from near the cave floor (see Ronen 1976:Fig. 1). Based on stratigraphic position, the Skhul hominids might fall into two

Group 2 Steinheim, Kabwe 1, Skhiil 4, Skhiil 9, Jebel Qafzeh 6, Gibraltar, La Chapelle 1, La Ferrassie 1, Jebel Irhoud 1. Group 3 Predmost 3, Predmost 4, Cro-Magnon 1, Mladec 1, Abri Pataud 1, Oberkassel 1. 77

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02

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Figure 4:3. Dendrogram from sum of squares cluster analysis with Skhiil 4, 5 and 9, Jebel Qafzeh 6 and 9 and Jebel Irhoud 1.

A2 0

2

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Figure 4:4. Correspondence analysis with Skhfil 4, 5 and 9, Jebel Qafzeh 6 and 9 and Jebel Irhoud 1.

78

in all of the analyses (Table 1:2). The placement of Skhiil 5 varied depending on the crania included and the number of variables used in the analysis. In general, it either clustered with anatomically modem Upper Palaeolithic crania from Europe, or with archaic Homo sapiens crania such as Steinheim, Petralona and Kabwe 1 (cf. Figs. 4:3, 4:4, 4:7, 4:8, 4:10, 4:11, 5:9, 5:10, Tables 4:2, 4:4, 5:4). Skhiil 5 also grouped with Jebel Qafzeh 9 (Figs. 4:3, 4:4, 4:5, 4:6, Tables 4:2, 4:3). On the analysis of the occipital squama (Fig. 5: 14) Skhfil 5 grouped with the anatomically modem crania. On the analysis of non-metric cranial traits (Fig. 5:15) Skhiil 5 was close to Saccopastore 1, and they are both in an intermediate position between archaic and anatomically modem crania. On the analysis of mandibular non-metric traits (Fig. 5: 16) the mandible of Skhiil 5 grouped with anatomically modem crania and was well separated from the Neanderthals from western Asia and Europe and the mandibles from Border Cave and Klasies River Mouth, South Africa. On the ordered similarity matrix (Table 4:7) Skhiil 5 was found to be similar to Jebel Qafzeh 9, Jebel Irhoud 1 and Amud I not very similar to Skhiil 4, Tabiin Cland Jebel Qafzeh 6. Where included in the same analyses, Skhiil 5 displays a closer relationship to Jebel Qafzeh 9 than to Jebel Qafzeh 6 (Figs. 4:3, 4:4, 4:5, 4:6, Tables 4:2, 4:7).

closer to the Neanderthal Tabiin Cl. It should be noted that the views detailed above were reached prior to the new dating of the Skhiil site and so were based on a younger dating of the Skhiil hominids. Thoma's ( 1957, 1958, 1965) conclusions are different to the results of the present study in that Skhiil 4 was aligned with the Neanderthals more so than Skhiil 9. This placement of Skhiil 4 reflects the sagittal contour of the cranium, but not with its overall morphological configuration, especially the non-metric features (i.e. the lack of Neanderthal traits). Coon (1963) saw similarities between Skhiil 4 and the Neanderthals, while Suzuki (1970b) found Skhiil 4 to be similar to Amud 1, but with the former being slightly more advanced. These two hominids (Skhiil 4 and Amud 1) were found to be similar on the ordered similarity matrix (Table 4:7). Suzuki (1970b:187) goes so far as to suggest "greater similarity between Skhiil IV and Amud I than between Skhiil IV and V", which is consistent with the ordered similarity matrix (Table 4:7). Skhiil 4 does have a long and relatively low cranium with some postbregmatic and pre-lambdoidal flattening which gives it a resemblance to Neanderthals. It does not however, have marked occipital angulation or protrusion or a Neanderthal occipitomastoid region in that there is no occipital torus or suprainiac fossa and the mastoid process is large. The frontal is also quite steep, especially the supraglabellar region. The face of Skhiil 4 displays marked alveolar prognathism, and not the mid-facial prognathism of the Neanderthals. The orbits are rectangular in shape and not circular, the malars are not inflated, and the superciliary arch is distinguishable from the supraorbital trigone on the prominent supraorbital region. The mandible of Skhiil 4 also displays some Neanderthal reminiscent features in that it has a slight retromolar gap and the mental foramen located under the first molar. However, the symphyseal region displays a distinct protrusion or chin.

Skhiil 9 initially clustered with Jebel Qafzeh 6, and then with other archaic Homo sapiens such as Steinheim, Jebel Irhoud 1 and Kabwe 1, and the Neanderthal Gibraltar 1 (Fig. 4:3, Table 4:2). On the correspondence analysis (Fig. 4:4) Skhiil 9 occupied an intermediate position between the archaic and anatomically modem crania and was well separated from Skhiil 5 and Jebel Qafzeh 9. Skhiil 4 and Jebel Qafzeh 6 were not markedly separated from Skhiil 9 (Fig. 4:4). There were high loadings on the variables that document the shape of the posterior cranium and the size of the face (Fig. 4:4, Table

A:8). In earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986) Skhiil 4 and 9 were found to group with Petralona and Kabwe 1, while Skhiil 5 clustered with Jebel Qafzeh 6.

The placements of Skhiil 5 and 9 do fit with their morphological configurations. Skhiil 5 displays little resemblance to the Neanderthals, but is close to both archaic Homo sapiens and fully anatomically modem crania as shown in the way that it is often referred to as a proto-CroMagnon (Howell 1957, 1958; Thoma 1965; Vandermeersch 1981, 1982). Whilst Skhiil 9 does have a long and low vault with some postbregmatic and pre-lambdoidal flattening on the parietals, it displays many similarities with other archaic Homo sapiens material such as Jebel Irhoud 1 and Kabwe 1. The more archaic nature of Skhiil 9 compared to Skhiil 5 may reflect its placement earlier within the stratigraphic sequence of Layer B. Rosenberg (1998) has indicated that Skhiil 9 has a relatively long pubis when compared to modem males, which aligns it more with the Neanderthals. On a bivariate plot of femoral head diameter (squared) against pubic length, Skhiil 9 was positioned with Tabiin C 1, Krapina 208, Kebara 2 and La Ferrassie 1, whereas Skhiil 4 was placed near Upper Palaeolithic material from CroMagnon and Predmost (Rosenberg 1998:Figure 3). Jebel Qafzeh 9 was well separated from the Skhiil material on this plot.

The results from this study are generally consistent with those of Corruccini (1974a), Howells (1970, 1989), Stringer (1974b, 1978, 1989) and Brauer and Rimbach (1990). The morphological affinities of the Skhiil material have been debated since their discovery. After their extensive study of the material, McCown and Keith (1939) concluded that the Mount Carmel population, within which they included the material from both the caves of Tabiin and Skhiil, were in the "throes of evolutionary change" diverging to produce both the Neanderthal and modem forms. Others such as Brace (1964) and Wolpoff (1980a) suggested that the Skhiil hominids were transitional between Neanderthals and anatomically modem types. Thoma (1957, 1958, 1965), following earlier suggestions by Keith and McCown (1937), Ashley-Montagu (1940) and Dobzhansky (1944), concluded that the Skhiil fossils, examined together or individually, demonstrated a morphological pattern of variation that was characteristic of "cross-breeds". That is, the Skhiil population was the result of hybridisation between Neanderthals and anatomically modem forms. Thoma found that Skhiil 4 and 5 were modem in overall configuration, whereas Skhiil 9 was

Howells ( 1970), using a multiple discriminant analysis based on modem crania, also found clear separation between Skhiil 5 and Neanderthal crania, with the former falling near the modem crania. In another analysis using principal 79

9 to be positioned closer to Neanderthals than Skhiil 5, and not far removed from European Upper Palaeolithic material.

components analysis Howells (1989:Figure 16) found Skhiil 5 to group with Jebel Irhoud 1, Kabwe 1 and then La Chapelle 1 and La Ferrassie 1. The group was well separated from the other component of the cluster analysis, which was made up of modem Homo sapiens samples and a group of "premodem " skulls including Jebel Qafzeh 6 (Howells 1989:Figure 16). Howells ' (1989) analysis indicates that Skhul 5 displays a less modem morphology than Jebel Qafzeh.

In earlier analyses Skhiil 1 was found to group with Skhiil 5 and Teshik-Tash (Habgood 1982, 1984b; Habgood and Walker 1986). Minugh-Purvis (1998) found Skhul 1 to be morphologically different from the Krapina 1 child that displays Neanderthal features, while Tillier ( 1998) found Skhiil 1 to be similar to Upper Palaeolithic children from Europe . Skhiil 1 displays numerous similarities with Jebel Qafzeh 11 when age and growth patterns are taken into consideration (Minugh-Purvis 1998).

In a study of the frontal squama, Hills and Brothwell (1974) found that Skhul 5 fell near an Upper Palaeolithic group . Using both bivariate plots and multivariate analyses Stringer (1978) found that Skhiil 5 was aligned with modem crania and was very similar to Jebe l Qafzeh 6 and 9. Stringer (1978) demonstrated that Skhiil 5 has a flat, non-Neanderthal face. Skhiil 9, on the other hand , displays a combination of archaic features , especially evident in the vault shape , and more modern features such as the form of the supraorbital region . On parietal indices String er (1978) found Skhiil 9 to fall with archaic and not modem hominids , whereas on other plots it fell with the more modem. material. On a multivariat e analysis of facial measurements , Turbon et al. ( 1997) found Skhul 5 to be similar to Jebel Qafzeh 6 and 9 and more similar to Upper Palaeolithic material than to western Asian and European Neanderthals , although Skhiil 5 was closer to the Neanderthals than the Jebel Qafzeh material.

Stringer et al. ( 1979) placed Skhiil in their Homo sapiens Grade 3b. Corruccini (1992) argued that the Skhiil crania as a group are intermediate between Upper Palaeolithic samples and Neanderthals and should not be referred to as "fully anatomicall y modern". This position would be consistent with Howell s ( 1989) results . Overall , the Skhiil sample displays a predominance of modem characteristics, but some of the individuals such as Skhiil 4 and 9 do preserve archaic sagittal contours , which give them a superficial resemblance to the Neanderthals and the mandible of Skhul 4 has a slight retromolar gap, mental foramen under the first molar and a symphyseal protrusion or chin. Skhul presents a variable hominid sample that may best be described as an early morphologically generalised sample with links to similar hominids from Africa such as those from Jebel Irhoud. As Howells has stated The Skhiil people must be diagnosed ...as archaic modem man, with a small number of Neanderthal-like details which are not general but distributed over different parts of different individuals. (1973a:93)

Corruccini (1992) pointed out that most studies only included Skhiil 5 and not Skhiil 4 and 9 and so underestimate the variation in the sample. Using size and shape distance statistics. In a pairwise clustering analysis Skhiil 5 grouped with Steinheim and Ehringsdorf and then joined with Zuttiyeh and the western Asian and European Neanderthals (Corruccini 1992:Fig. 2) . Skhiil 4 and 9 grouped together and then joined a group including Jebel Irhoud 1 and 2, Jebel Qafzeh 6, Florisbad and Omo 2 before becoming part of the bigger cluster including Skhiil 5 and the Neanderthals (Corruccini 1992:Fig. 2). The last group to join the cluster were a group of eastern European Upper Palaeolithic crania . When linear generalised distances were calculated different relationships emerged . Skhiil 5 adopted a neutral position by being as similar to the Neanderthals as to the eastern European Upper Palaeolithic crania, whereas Skhiil 4 and 9 were closer to the Neanderthals . Corruccini 's study highlights the morphological variation among the Skhiil hominids and aligns them with archaic crania from Africa (what he calls Presapiens) and not with anatomically modern Upper Palaeolithic crania. The relative positions of Skhiil 4 and 5 in Corruccini ' s ( 1992) pairwise clustering analysis were different to the present investigation where Skhiil 4 was aligned with the Neanderthals and Skhul 5 with Upper Palaeolithic material. These relative positions seem to better match the overall morphology of the Skhiil materia l.

J ebel Qafzeh-J ebel Kafzeh The remains of at least 15 individuals were recovered from Levalloiso-Mousterian levels at Qafzeh Cave, Israel (Fig. 4 :1; Neuville 1951; Oakley et al. 1975; Vandermeersch 1981). Additional hominid material was found in Upper Palaeolithic levels (Oakley et al. 1975). The Middle Palaeolithic hominids predominantly came from Layer E of the original excavations and Layer XVII of the more recent excavations undertaken at the site (Oakley et al. 1975; Vandermeersch 1981). The hominid remains from Jebel Qafzeh are clustered near the mouth of the cave within Layer XVII (Hovers et al. 1995:Fig. 1), which has been interpreted as a deliberate burial tradition at the site. It is also worth noting that associated with the Jebel Qafzeh 11 individual was a set of fallow dear antlers that may be an example of a burial gift. Gargett (1999) has questioned these interpretations and proposed that the Jebel Qafzeh "intentional burials " were actually the result of crushing and natural burial from a rockfall , possibly caused by tectonic activity.

Brauer and Rimbach (1990) found on a principal components analysis using facial measurements that Skhul 5 was placed near Jebel Qafzeh 6 and among European Upper Palaeolithic and late Pleistocene northern African material , whereas when facial and vault measurements were used Jebel Qafzeh 6 was again placed with late Pleistocene northern African material while Skhiil 5 was somewhat removed but still closer to the late Pleist ocene northern African material, especially Gebel Sahaba. Brauer and Rimbach ( 1990) also found Skhiil 4 and

The two most complete adult individuals, Qafzeh 6 and 9, and the most complete immature cranium, Qafzeh 11, will be dealt with here, but descriptions of much of the other 80

prognathism, the orbits are large and sub-rectangular in shape and the nasal aperture is moderately large (Tillier 1984). The mandible displays a prominent chin, ascending rami that do not angle backwards markedly, single mental foramina positioned between the premolars, and the second molars are partly obscured by the ascending rami.

material may be found in Arensburg and Tillier (1983), Tillier (1979), Tillier and Vandermeersch (1976), Vandermeersch and Tillier ( 1977), and Vandermeersch ( 1981). The postcranial material from Qafzeh is robust, but essentially modem in form and displays similarities with the Skhul sample and contrasts with western Asian Neanderthals (Trinkaus 1983b, 1993, 1995; Trinkaus et al. 1998; Vandermeersch 1981). As with the Skhiil sample, the Jebel Qafzeh individuals can be described as possessing linear bodies with narrow trunks and long robust limbs (Trinkaus 1995 and references therein).

Establishing an absolute chronology for Jebel Qafzeh has been difficult. Initial assessment of the Jebel Qafzeh lithics (Neuville 1951), and analyses of microfauna and sedimentology (Bar-Yosef 1989, 1998; Bar-Yosef and Vandermeersch 1981; Farrand 1979; Jelinek et al. 1973; Tchernov 1981, 1998; Vandermeersch 1981a) suggested an age of 70,000 and 80,000 years BP for the Middle Palaeolithic hominid bearing levels at Jebel Qafzeh. A major faunal difference between the sites of Tabiin and Jebel Qafzeh is the absence, in Tabun Layers C and D of two archaic rodents (Mastomys batei and Avicanthis ectos) that are present in Tabiin Layers E and F (the latter only) and at Jebel Qafzeh (Bar-Yosef 1989; Bar-Yosef and Vandermeersch 1981; Tchernov 1981). This pattern suggests the possible contemporaneity between Jebel Qafzeh and Tabiin Layers E and F. It was suggested that these differences may be due to the varying habitats of the two sites, with Qafzeh being closer to the refuge of the Jordan Valley and so it has later examples (Jelinek, A.J. 1982a). However, other sites, such as Amud Cave, which are also close to the Jordan Valley do not have any remains of these two rodents (Bar-Yosef 1989). The presence of an archaic form of the rodent Myomimus roachi (qafzehsis) is also suggested as demonstrating an early date for Qafzeh (BarYosef 1989; Bar-Yosef et al. 1986). It has also been argued that there is a correlation of a depositional hiatus between Layers D and C at Tabiin with an erosional event of unknown duration following the Mousterian levels at J ebel Qafzeh (Bar-Yosef and Vandermeersch 1981). If correct, this correlation would equate the Jebel Qafzeh Middle Palaeolithic levels with Tabiin Layer D, but the two events could reflect local climatic and/or karstic events and so are not related (Farrand 1979; Jelinek, A.J. 1982b).

Jebel Qafzeh 6 (Vallois and Vandermeersch 1975; Vandermeersch 1981) is an adult cranium with a moderately high and well-rounded vault and little postorbital constriction. The supraglabellar region is steep, while the remainder of the frontal gently recedes. Parietal bossing is quite marked. There is a certain degree of pre-lambdoidal flattening, but the occipital is generally full and rounded, with none of the typical Neanderthal features (Santa Luca 1978). The preserved mastoid process is quite large and broad, and does not taper. The supraorbital torus is welldeveloped, but can be separated into medial and lateral segments. The broad, flat face is essentially orthognathic, with only slight alveolar projection. The sub-rectangular orbits are quite large, while the rounded nasal aperture is not very high. The nasal bones project moderately, and the maxillae display distinct canine fossae. Jebel Qafzeh 9 is a young adult skeleton (Plates 5 and 6; Vandermeersch 1981). The cranium is higher and narrower than Jebel Qafzeh 6, with less markedly developed supraorbital arches that are more clearly divisible into medial and lateral segments. The high rounded vault displays only slight parietal bossing and no lambdoidal flattening. The occipital does not display an occipital torus or suprainiac fossa. The remaining mastoid process is quite large and does not taper. There is no occipitomastoid crest. The relatively flat face is narrower than that of Jebel Qafzeh 6 with smaller sub-rectangular orbits and a larger nasal aperture. The marked alveolar projection of Jebel Qafzeh 9 is greater than that of Jebel Qafzeh 6. The mandible of Jebel Qafzeh 9 (Plate 8) has a relatively short and narrow ramus, a well-developed mental eminence and no retromolar gap. The condylar and coronoid processes are of similar size and the mandibular notch is deep with the maximum depth towards the centre of the notch. The Jebel Qafzeh 9 mandible, along with Skhiil 4 and 5, does not have the horizontal-oval mandibular foramen form.

Typological analysis of the lithic material from Jebel Qafzeh suggested it was similar to the material from the "chimney" deposits at Tabiin and Skhiil Level B (Jelinek, A.J. 1982a, 1982b), which does not equate with the analyses of microfauna and sedimentology. This typological analysis approach does not consider the influence of raw material on flake form and the applicability of the patterns established for sites in the Wadi el-Mughara outside of the Mount Carmel region has not been established (A.J. Jelinek pers. comm.). Other studies have identified similarities with Tabiin C-type (Bar-Y osef 1992, 1998).

Jebel Qafzeh 11 is an immature skeleton given an age of approximately 12-13 years old (Tillier 1984). The vault, which displays some postmortem crushing on the right side, is high and well-rounded, with little brow-ridge development and no lambdoidal flattening or occipital bunning. The supraglabellar segment of the frontal is especially steep. There is little postorbital constriction. The undistorted left parietal is relatively vertical and so the maximum cranial breadth would have been high up on the parietals. Jebel Qafzeh 11 has small mastoid processes that taper and a rounded foramen magnum. There is no occipital torus, suprainiac fossa or marked occipitomastoid crests. The maxillae are detached from the vault. On the reconstruction (the mandible has been used to help position the face) the face is relatively short and displays only moderate

Revised aspartic acid racemisation dates were obtained from Jebel Qafzeh (Masters 1982), but there are problems with because there are no radiometric dates from the site for calibration, the hominid material used had been treated with preservatives that could cause discrepancies, and because a faunal sample from an Upper Palaeolithic level provided a date older than the hominid samples from Mousterian levels. The Upper Palaeolithic faunal sample produced revised dates of 47,000 and 54,000 years BP, while the hominid material, Jebel Qafzeh 3 and 6, provided revised dates of 39,000 and 45,000 years BP and 40,000 and 46,000 years BP respectively, and two faunal samples from Mousterian levels 81

breadth of the frontal squama, and the sagittal contour of the posterior cranium (Fig. 4:4, Table A:8). On the analyses of cranial (Fig. 5: 15), and mandibular non-metric traits (Fig. 5:16) Jebel Qafzeh 9 also fell with the anatomically modem Upper Palaeolithic crania. The immature Jebel Qafzeh 11 grouped with modem crania, but was separated from the other Jebel Qafzeh crania and the Skhul material (Figs. 4:5, 4 :6, Table 4:3). There were high loadings on the variables that describe nasal form, sagittal curvature and breadth of the frontal and occipital bones (Fig. 4:6 , Table A:9).

below the hominid samples tested gave revised dates of 59,000 and 68,000 years BP and 68,000 and 78,000 years BP (Masters 1982). It is of interest to note that the revised aspartic acid racemisation dates for the Jebel Qafzeh hominids are similar to the revised aspartic acid racemisation dates from Skhul (Masters 1982). As with the site of Skhul, much older dates for the Mousterian levels at Jebel Qafzeh have been obtained by the more recent use of radiometric methods including electron spin resonance (ESR) and thermoluminescence (TL).

Table 4:3. K-means cluster analysis with Skhfil 4, Jebel Irhoud 1 and Jebel Qafzeh 6, 9 and 11. Analysis includes 75.2% of the total within-group variance.

Twenty burnt flints from Layers XVII-XXIII at Jebel Qafzeh provided TL dates ranging between 85- 110,000 years BP with a weighted mean TL age of 92,000+/-5,000 years BP (Valladas et al. 1988). There was no systematic increase in age with depth of flint sample used .

Group 1 Steinheim, Kabwe 1, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Jebel Irhoud 1, Skhul 4, Jebel Qafzeh 6.

Six large mammal teeth from Layers XV -XXI have produced 14 ESR age determination s with no systematic increase with depth, as with the TL results . The samples produced an average age of 96,000+/-13, 000 years BP for early uptake (EU) and 115,000+/- 15,000 years BP for linear uptake (LU)(Schwarcz et al. 1988). Griin (Griin and Stringer 1991) added an additional sample and recalculated the average ages resulting in figures of 100,000+/- 10,000 years BP for EU and 120,000+/-8,000 years BP for LU. The EU ESR average age is consistent with the TL results, whereas the LU ESR average age is older.

Group 2 Predmost 3, Predmost 4, Cro-Magnon 1, Mladec 1, Abri Pataud 1, Oberkassel 1, Jebel Qafzeh 11. Group 3 Skhfil 5, Combe Capelle, Chancelade , Dolni Vestonice 3, Oberkassel 2, Jebel Qafzeh 9.

These results are comparable to those of Corruccini (1974a), and Stringer (1974b, 1978), Brauer and Rimbach (1990).and are compatible with earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986).

McDermott et al. (1993) used the mass-spectrometric uranium-series (U-series) method to provide ages for two bovid tooth samples from Layer XIX analysed by Schwarcz et al. (1988). Sample 368DE provided an age of 106,400+/2,400 years BP, which is close to the EU ESR age of 105,000+/-2,000 years BP (McDermott et al. 1993). Sample 371EN provided an age of 88,600+/-3,200 years BP, which was much younger than the EU ESR age of 103,000+/19,000 years BP (McDermott et al. 1993). It should be noted that the EU method provides younger age estimates than the LU method and so these U-series results , especially for sample 371EN, would be considerable younger than their LU results (371EN-125 ,000+/ -22,000 and 368DE- 115,000+/-800 years BP).

Howells (1989:Figure 16) found Jebel Qafzeh 6 to cluster with "prehistoric modem" crania , including Mladec 1, Fish Hoek and Keilor, and then join a large grouping of modem and "prehistoric modem" crania. This group was well separated from the other component of the cluster analysis, which was made up of Skhul 5, Jebel Irhoud 1, Kabwe 1 and then La Chapelle 1 and La Ferrassie 1 (Howells 1989:Figure 16). Howells' ( 1989) analysis indicates that Jebel Qafzeh 6 displays a more modem morphology than Skhiil 5. Corruccini ( 1992) also achieved a separation of Skhiil 5 and Jebel Qafzeh 6. on a pairwise clustering analysis. However, Jebel Qafzeh 6 did not cluster with anatomically modem material , but with archaic crania such as Jebel Irhoud 1 and 2, Omo 2, Florisbad and Skhiil 4 and 9 (Corruccini 1992:Fig. 2). Skhiil 5 grouped with Steinheim and Ehringsdorf and then joined with Zuttiyeh and western Asian and European Neanderthals (Corruccini 1992:Fig . 2).

The TL, ESR and U-series results indicate that the Middle Palaeolithic layers at Jebel Qafzeh accumulated over a relatively short time period . The TL, ESR and U-series determinations and the analyses of microfauna and sedimentology suggest an oxygen isotope stage 5 age for the Middle Palaeolithic levels at J ebel Qafzeh, with the hominids probably dated at around 90-100 ,000 years BP (Table 5:1; Fig. 4.4).

In an analysis of the frontal, Simmons ( 1991) also found a strong similarity between Jebel Irhoud 1 and the Jebel Qafzeh hominids , especially with respect to the browridge area. In another analysis of the frontal bone, Simmons et al. (1991 :Figure 1, 2 and 4) found Jebel Qafzeh 6 to be positioned with the material from Skhiil on bivariate plots from the principal components analyses of log size and shape and log shape and the UPGMA cluster analysis. Not far removed were Zuttiyeh , Tabiin Cl and Amud 1. Jebel Qafzeh 9 was separated from Jebel Qafzeh 6 and the Skhiil material and grouped with the Upper Palaeolithic associated Jebel Qafzeh 1 and 2. This study suggests that Jebel Qafzeh 9 displays a more modem morphology than Jebel Qafzeh 6 and Skhiil 4, 5 and 9. The current analyses also found Jebel

The nature of the Levalloiso -Mousterian stone tools, radiometric dating results and the morphology of the hominid remains from Jebel Qafzeh suggest a date equivalent to, or slightly younger than , the material from Skhfil. Jebel Qafzeh 6 was found to group with archaic crania such as Skhiil 9, Steinheim, Kabwe 1, and Jebel Irhoud 1 (Figs. 4:3, 4:4, 4:5, 4:6, Table 4 :2). Jebel Qafzeh 9 clustered with Skhiil 5, and anatomically modem Upper Palaeolithic crania from Europe (Figs. 4:3, 4:4 , 4 :5, 4:6, Tables 4:2, 4:3) . There were high loadings on the variables that document the 82

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83

the chimney deposits in the inner cave and on a terrace at the exterior edge of the outer cave and subdivided the Tabun deposit into Layers A to G. A.J. Jelinek's (Jelinek et al. 1973) excavations concentrated on the intermediate chamber and the stepped vertical profile at the entrance of the inner cave left by Garrod. The levels identified by Jelinek (1982a; Jelinek et al. 1973) can generally be equated with those identified by Garrod: Unit I, beds 1-16 Layer C [LevalloisoMousterian] Unit I, beds 17-26 Layer C [LevalloisoMousterian] Units II-V = a solution cavity between Layers DIC Unit IX = Layer D [Levalloiso-Mousterian] Unit X = Layer Ea-D Unit XI = Layer Eb-Ea [Amudian/ AcheuleoYabrudian] Unit XII= Layer Ee-Eb [Acheuleo-Yabrudian] Unit XIII = Layer Ed [Y abrudian] Unit XIV = Layer G [Late Acheulian]

Qafzeh 9 to group with more modern crania, but also to be placed near Skhul 5. Turb6n et al. (1997) found Jebel Qafzeh 6 and 9 to be similar to Skhul 5 and more similar to Upper Palaeolithic material than to western Asian and European Neanderthals . Brauer and Rimbach (1990) found on a principal components analysis using facial measurements that Jebel Qafzeh 6 was placed near Skhul 5 and among European Upper Palaeolithic and late Pleistocene northern African material, whereas when facial and vault measurements were used Jebel Qafzeh 6 was placed with late Pleistocene northern African material while Skhul 5 was somewhat removed but still closer to the late Pleistocene northern African material , especially Gebel Sahaba. On a bivariate plot of femoral head diameter ( squared) against pubic length , Jebel Qafzeh 9 was placed near Upper Palaeolithic material and modem males and was well separated from the Skhul 4 and Skhul 9 and Neanderthal material -Tabiin Cl, Krapina 208, Kebara 2 and La Ferrassie 1 (Rosenberg 1998:Figure 3).

The depositional units (Layers A to G) identified by Garrod and Bate (1937) are still useful for general discussions of the site 's stratigraphy.

The results from the present study are consistent with the morphology of the three Jebel Qafzeh crania . Jebel Qafzeh 9 appears more modem than J ebel Qafzeh 6 in that it has a higher cranial vault, lacks lambdoidal flattening and has a reduced supraorbital region . The two crania are also similar to the Skhul material, especially Skhul 5, in overall cranial morphology (Stringer 1978; Vandermeersch 1981a). Jebel Qafzeh 6 and Skhul 5 are very similar in their sagittal contours (Vandermeersch 1981a). Jebel Qafzeh 11 has a cranial morphology that also displays numerous similarities to Skhul 1 when age and growth patterns are taken into consideration (Minugh-Purvis 1998). Schwartz and Tattersall (2000) found Jebel Qafzeh 11 and possibly Jebel Qafzeh 8 and 9 to possess the inverted "T" and mental fossae chin morphology found on Upper Palaeolithic and modem mandibles.

Tabun Cl (Plate 5; McCown and Keith 1939; Trinkaus 1984a) is the skeleton of an adult female . The relatively small cranium has a low vault, a rounded occipital and marked postorbital constr1ct1on. The well-developed supraorbital torus does not taper laterally. The fragmentary facial skeleton of Tabun Cl, which was reconstructed using the mandible to locate the position of the maxillary dentition, displays only moderate mid-facial prognathism . This is supported by the relatively high nasiofacial angle and the large ramus index of the mandible (Trinkaus 1984a). There are "classic" Neanderthal features (Table 5:6) evident on the facial skeleton of Tabun Cl including a large, prominent nasal aperture and round orbits. Apart from large occipitomastoid crests, the posterior cranium of Tabun Cl seems to lack most of the "classic" Neanderthal features (Table 5:6), such as a lack of lambdoidal flattening, occipital bunning, a marked occipital torus, and a suprainiac fossa (contra Santa Luca 1978), but this may be due to problems of preservation and reconstruction.

Stringer et al. (1979) placed the Jebel Qafzeh hominids, along with the Skhul material, in their Homo sapiens Grade 3b.

The mandible of Tabun Cl displays a receding symphysial region, a retromalar space and the mental foramina positioned under the first molar. The coronoid processes would have been large and the mandibular notch is shallow with the deepest point near the condyle. All of these Neanderthal features (Table 5:6) suggest facial prognathism. Tabun Cl does lack the "classic" Neanderthal horizontaloval mandibular foramen pattern (Smith 1978; Trinkaus 1983a).

Mugharet Et-Tabiin Hominid remains have been recovered from Layers E, C and B within the long stratigraphic sequence at et-Tabun cave in the Wadi el-Mughara, Mount Carmel, Israel (Fig. 4:1 ; Garrod and Bate 1937; McCown and Keith 1939; Oakley et al. 1975). The cave of et-Tabiin consists of three chambers the outer , inner with chimney and intermediate. Much of the hominid material is fragmentary and so only the Tabun Cl skeleton and the Tabun C2 mandible will be dealt with here. Tabun Cl is generally regarded as an intentional burial (McCown and Keith 1939 ; Oakley et al. 1975), but Gargett ( 1989) has suggested that the location of Tabun C 1 near the back wall of the cave and below an overhang would have protected the body from disturbance while natural burial occurred .

The postcranial remains of Tabun Cl also display Neanderthal features such as the morphology of axillary border of the scapula and low crural indices, but there is less hypertrophy and relatively higher brachial indices (Table 4:8; McCown and Keith 1939; Trinkaus 1981, 1984a). Other postcranial material from Layer C also displays Neanderthal morphological similarities (McCown and Keith 1939; Stefan and Trinkaus 1998).

Two major excavations have been undertaken at et-Tabun cave. The excavation of Garrod and Bate (1937) focused on 84

Tabiin C2 is a large and robust adult mandible (McCown and Keith 1939; Quam and Smith 1998; Rak 1998; Trinkaus 1983a, 1984a). The large anterior teeth display marked dental attrition. Large single mental foramina are situated below the anterior edge of the first molar. The rami are moderately broad, but appear narrower because of the presence of marked pre-angular notches that accentuate the retromolar spaces. The condylar and coronoid processes are of similar size and the mandibular notch is deep with the maximum depth towards the centre of the notch. Rak (1998) has argued that Tabun C2 only has retromolar spaces because of the marked pre-angular notches and that this morphology is different to the Neanderthal pattern where the retromolar space is caused by more anteriorly placed third molars.

Upper Pleistocene in age and that the sediments of the earlier layers were predominantly fine-grained sand because the sea was very near the cave, whereas the sediments of Layer D were siltier because the sea had retreated. There is an erosional discontinuity between Layers D and Layer C and so there may be a significant time interval between the two layers, enough time for a solution cavity that had formed in the cave to be filled (Farrand 1979; Jelinek, A.J. 1982a). The sediments of Layer C suggested that an opening in the cave roof had formed, allowing material from the plateau above to enter the cave (Farrand 1979). Layers C and B may have been deposited in a relatively quick time interval. There is a major fauna} change between Layers C and B, a dominance of gazelle to one of Dama (Garrod and Bate 1937), which may be related to the opening of this hole (chimney) in the cave roof, and a change in cave use from an occupation site to a specialised butchering station of Dama that had been driven into the cave from the plateau via the chimney (Farrand 1979; Jelinek, A.J. 1982a, 1982b).

Brothwell ( 1961: 156) observed that the exact form of the symphyseal area of Tabiin C2 is "obscured by breakage and loss". This caution has generally been ignored in assessments of chin development on Tabiin C2. Even when the damage is taken into consideration, the symphyseal region on Tabon C2 does display some anterior alveolar prognathism caused by the roots of the incisors and has what I have assessed as slight development of a mental eminence. Quam and Smith ( 1998) observed that whilst a mental trigone was probably present on Tabiin C2, the preserved morphology of the symphysial region precludes a definite identification. They acknowledged that the morphology of the symphysial region was more comparable to J ebel Qafzeh 9 and Skhul 4 and 5 than to Amud 1, Kebara 2 and Tabiin Cl. Tabiin C2 does not have the chin morphology found on the mandibles of Jebel Qafzeh 11, 8 and 9 or Upper Palaeolithic and modern individuals and differs from the configuration on the Skhul mandibles (Schwartz and Tattersall 2000). Unlike the Tabiin Cl mandible, Tabon C2 has the horizontal-oval mandibular foramen pattern (Smith 1978; Trinkaus 1983a). The features of Tabun C2 suggest that the cranium that it belonged to would have displayed only moderate facial prognathism.

Two main dating sequences for the sedimentary units at Tabiin have been proposed (Bar-Yosef 1989, 1992; Farrand 1979; Jelinek, A.J. 1982a, 1982b). Farrand (1979) and A.J. Jelinek (1982a, 1982b) argued that the beginning of the sedimentary record at Tabiin could be equated with a major warm period during the last interglacial, oxygen isotope stage Se, and dated at approximately 120,000 years BP. Layer E was dated to between 110-80,000 years BP. Layer D was placed in oxygen isotope stage 4 and a relatively short erosional disconformity separated these deposits from Layer C. Layers C and B were equated with oxygen isotope stage 3, with Layer C being dated to around 50,000 years BP (Farrand 1979; Jelinek, A.J. 1982a, 1982b). Jelinek has revised this chronology based on thermoluminescence dating and a reallocation of the large sand dune near Tabun from oxygen isotope stage 5e to oxygen isotope stage 7, which gives the site a much earlier date and a longer depositional duration (Table 5: 1; Mercier et al. 1995).

The larger and more robust Tabon C2 mandible differs from the mandible of Tabiin C 1 in many features with the two mandibles displaying marked morphological variation, which may be related to temporal differences, sexual dimorphism, individual variation within a population, or even different morphological types.

studying faunal changes in the Mediterranean area, Higgs ( 1961; Higgs and Broth well 1961) suggested Skhul was as much as 10,000 years later than Tabun, a separation that had earlier been proposed by Howell (1958).

Bar-Yosef (1989, 1992) proposed a more extended sequence for Tabiin based on biostratigraphy, geomorphology and radiometric dates from other Levantine sites. Layers G and F were equated with oxygen isotope stage 7 and oxygen isotope stage 6 respectively, Layer E to oxygen isotope stage 5/6 and Layer D to oxygen isotope stage 5 and dating to around 100,000 years BP. Bar-Yosef (1989) proposed a relatively long erosional phase between Layers D and C, with Layers C and B being placed in oxygen isotope stage 4 and early oxygen isotope stage 3 and possibly dating to less than 60,000 years BP (Layer C could be as old as 90,000 years BP if the erosional phase was short [Bar-Yosef 1992]). Both schemes have the deposits ending at around 40,000 years BP. Mercier et al. (1995) have questioned the data that Bar-Yosef used to formulate his chronology, especially the radiometric dating results.

The sediments at Tabiin can be divided into three major depositional units - a lower fine-grained sandy deposit consisting of Layers G, F and E (upper Layer E has increasing amounts of silt), a middle loess deposit comprising Layer D and an upper red earth (washed-in terra rossa) deposit consisting of Layers C and B (Farrand 1979). Farrand (1979) suggested that the entire Tabun sequence was

Typological analysis of the stone tools from Tabiin by A.J. Jelinek suggested a continuous sequence of development from the Upper Acheulean to the late Mousterian, as demonstrated by a clear progression of a width-to-thickness ratio of flakes through the sequence (flakes become thinner with respect to their width) and a gradual increase in Levallois elements (Jelinek, A.J. 1975, 1977, 1981, 1982a,

A number of dating schemes have been proposed for the Tabiin sequence, but each method seems to contradict the others (see Bar-Yosef 1989, 1998; Griin et al. 1991; Jelinek et al. 1973; Schwarcz and Rink 1998; Tchernov, 1998; Valladas et al. 1998). Garrod and Bate ( 1937) concluded that Tabon Layers B and C were contemporary with Skhul Layer B. However, after

85

averaged results were obtained for early uptake (EU) and linear uptake (LU): Layer B EU 86,000+/-11,000 years BP LU 103,000+/- 16,000 years BP Layer C EU 102,000+/-l 7,000 years BP LU 119,000+/-1 l ,OOOyears BP Layer D EU 122,000+/-20,000 years BP LU 166,000+/-20 ,000 years BP Layer Ea EU 154,000+/-34,000 years BP LU 188,000+/-31 ,000 years BP Layer Eb EU 151,000+/-21 ,000 years BP LU 168,000+/- 15,000 years BP Layer Ee EU 176,000+/-10 ,000 years BP LU 199,000+/-7,000 years BP Layer Ed EU 182,000+/-3 1,000 years BP LU 213,000 +/-46,000 years BP It is worth noting that the results for Layers Ea and Eb produced stratigraphically reversed ages.

1982b). Using the width-to-thickness ratio of flakes A.J . Jelinek (1982a, 1982b) concluded that Skhul Layer Band the lowest layers at Qafzeh (L-M of Neuville [1951] and XVIXXIV of Vandermeersch [1966]) were younger than Tabun Layers D and C, and most probably equate to the "chimney " deposits at Tabun . This sequence is not consistent with radiometric dating results and faunal studies carried out on material from Tabun and other sites including Jebel Qafzeh and Skhul . From studies of the microfauna at Tabun , Tchemov (1981) proposed that Layer E be placed into the Riss-Wiirm ( early oxygen isotope stage 5), Layer D to the Riss-Wurm/Wurm transition (oxygen isotope stage 5/4 transition) and Layers C and B to the Wurm (oxygen isotope stages 4 and 3). Layer XVII at Jebel Qafzeh was equated with Layer D at Tabun and Layers C and B were equated with the hominid bearing levels at Kebara {Tchemov 1981:Fig. 10). This sequence would place the Jebel Qafzeh hominids earlier than most of the materia l from Tabun, includin g Tabun Cl and C2. The thermoluminescenc e dates of approximately 92,000 years BP from the hominid bearin g levels at J ebel Qafzeh (Vallado s et al. 1988) support this sequence. Tchemov (1998 :Figure 2) revised the chronology and placed Layer B in oxygen isotope stage 3-4, Layer C in oxygen isotope stage 5, Layer D in oxygen isotope stage 5- 6 and Layer E in oxygen isotope stage 7.

The result s would plac e Layer B into late oxygen isotope stage 5 or early oxygen isotope stage 4, Layer C into oxygen isotope stage 5, Layer D into late oxygen isotope stage 5 or early oxygen isotope stage 6 and Layer E into oxygen isotope stage 6 (Griin et al. 1991). These results are significantly earlier than the other proposed chronologies for the Tabun deposits and would place the Layer C hominids at older than 100,000 years BP, although the stratigraphic placements of the teeth used cannot be related to the hominids other than on a broad same layer basis. The ESR dates for Layer C are comparable to, although slightly older than, the average ESR dates of 81,000+l-15,000 years BP (EU) and 101,000+/-12,000 years BP (LU) for Layer B at Skhul (Stringer et al. 1989) .

The following radiocarbon dates have been obtained for Tabiin (Bar-Yosef 1989:Table 30.1; Vogel and Waterbolk 1963): Layer B 39,700+/-800 years BP (GrN-2534) Layer C 40 ,900+/ - 1000 years BP (GrN-2729) 51,000+4 ,800-3,000 years BP (GrN-7409) Unit I >47 ,000 years BP (GrN-7408) 51,000+4,800-3,800 (GrN-7409) 45 ,800+2,100-1,600 years BP (GrN-7410) Unit V -Bed 42 38,800+/-2,400 years BP (LJ-2084)

The mass-spectrometric uranium -series (U-series) method was used to provide ages for bovid tooth samples used by Griin et al. ( 1991) for the ESR determinations detailed above (McDermott et al. 1993). The method provided the following results : Layer B 50,690+/-230 years BP Layer C 94,840+430/-420 years BP 101,690+ 1,360/-1,340 years BP 105,360+ 2,580/-2,520 years BP Layer D 110,680+8801-870 years BP Layer Ea 159,100+/- 1,300 years BP 168,100+/-2,600 years BP

These dates should be regarded as minimum estimates as most of the deposits are probably beyond the practical limits of the method. Samples from the same layers were also found to contain some contamination , which may also have contributed to the young dates obtained (Jelinek et al. 1973). Revised aspartic acid racemisation dates of 63,000 and 65,000 years BP, and 68,000 and 71,000 years BP have been obtained from two bone samples from lower Layer C, but the bone sample used to provide the latter date may have been heated which would produce an "erroneously elevated date" (Masters 1982:5 I). A piece of unidentified bone that was said to have been associated with Tabun Cl (although C.B. Stringer pers. comm. suggested that the bone sample came from the Tabun C I skeleton) provided amino acid racemisation dates of 51,000 and 53,000 years BP (Masters 1982). A sample from the upper part of Layer E has produced revised asparti c acid racemisation ages of 83,000 and 87,000 years BP (Masters 1982). There are, however , problems with amino acid racemisation dating (Hare 1980).

These results are closer to, although still younger than, the ESR EU results on the same samples (Griin et al. 1991). Cervid (red dear) mandibular teeth from Layer E have also provided ESR dates of 215,000+/-22 ,000 years BP for EU and 290,000+/ -36,000 years BP for LU (Schwarcz and Rink 1998), which are older than the dates obtained by Griin et al. ( 1991). Mercier et al. (1995) undertook thermoluminescence (TL) dating on burnt flints from Jelinek ' s excavations at Tabun . The burnt flints had clear stratigraphical placement within beds and depositional units from Jelinek' s excavations. The following averaged TL ages were obtained (Mercier et al. 1995): 171,000+/-l 7,000 years BP Unit I-Layer C 212,000+/-22,000 years BP Unit II-Layer DIC 244,000+/-28,000 years BP Unit V-Layer DIC

Radiometric dating at the site has produced conflicting results . Twenty bovid teeth recovered during Garrod and Bates excavations at Tabun have been used for electron spin resonance (ESR) dating {Griin et al. 1991). The following 86

Unit Unit Unit Unit Unit

IX-Layer D X-Layer Ea-D XI-Layer Eb-Ea XII-Layer Ec(?)-Eb XIII-Layer Ed

263,000+/-27,000 270,000+/-22,000 306,000+/-33,000 350,000+/-33,000 331,000+/-30,000

out that towards the west wall, the area where the burial was recorded, Layer D becomes red, and that the burial may have come from this layer. It is of interest that Farrand stated that bones are preserved in Tabiin C and B, but not in beds D to G where bone material has been dissolved and redeposited as collophane and other secondary phosphate minerals. (1979:376) Whilst not conclusive or typical of the entire cave deposit, this observation would argue against the possibility that Tabiin Cl came from Layer D. It must also be remembered that there are other hominid remains from Layer C, some of which may have actually belonged to the Tabiin C 1 individual (Oakley et al. 1975; Quam and Smith 1998).

years BP years BP years BP years BP years BP

These results confirmed the long gap between Units I and II or Layers D and C. There is a stratigraphic inversion with the average ages for Units XII and XIII, but Mercier et al. ( 1995) suggested that Units XI, XII and XIII be grouped together. Mercier et al. ( 1995) performed isochron analysis on five burnt flints from Unit XI that yielded an average age of 287,000+/-20,000 years BP. They concluded that this analysis verified the antiquity of the deposits because it was in agreement with the TL average age of 306,000+/-33,000 years BP for Unit XI (Mercier et al. 1995).

The position of Tabiin Cl is especially important for establishing its relationship to the morphologically different Tabiin C2 mandible, which could reflect sexual dimorphism, individual vananon within a population, temporal differences, or even different morphological types. Tabiin C2 is from Layer C and most likely lower in the stratigraphy than Tabiin C 1 and therefore probably older.

These TL results are significantly older than the ESR average ages obtained by Grun et al. (1991) and the U-series results achieved by McDermott et al. ( 1993), but are comparable to the ESR LU date for Layer E reported by Schwarcz and Rink (1998). Mercier et al. (1995) suggested that the ESR ages were minimum estimates. These very early TL dates are also not compatible with the TL dates from Jebel Qafzeh where a Tabiin C-type industry is dated to 92,000 years BP as opposed to 170,000 years BP, or the averaged ESR results from Skhiil of 81,000+/-15,000 years BP (EU) and 101,000+/-12,000 years BP (LU). The dates also do not sit comfortably with the geomorphological or biostratigraphical analyses undertaken at Tabiin.

Tabiin Cl was found to group, initially, with the Neanderthals from Europe, and then with the Neanderthals from western Asia (Figs. 4:7, 4:8, Table 4:4). There were high loadings on the variables that describe the form of the posterior cranium and the size of the face (Fig. 4:8, Table Table 4:4. K-means cluster analysis with Tabiin Cl , Shanidar 1 and Amud 1. Analysis includes 67% of the total within-group variance.

Bar-Yosef. (1989:587) observed " ... there are many ambiguities in the chronological interpretations of the Tabiin sequence". The more recent radiometric dating attempts have not clarified the situation because the various dating methods used have provided different results and chronological schemes. A conservative dating scheme will be adopted here: Layer B 40-60,000 years BP Layer C 60-110,000 years BP Layer D 120-150,000+ years BP Layer E 150-250,000+ years BP

Group 1 Steinheim, Kabwe 1, Skhiil 5, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Amud 1, Shanidar 1, Tabiin Cl. Group 2 Predmost 3, Predmost 4, Cro-Magnon 1, Combe Capelle, Chancelade, Dolni Vestonice 3, Mladec 1, Abri Pataud 1, Oberkassel 1, Oberkassel 2.

This dating scenario would make the Tabiin Layer C hominids contemporary or slightly younger than the hominid material from Skhiil and Jebel Qafzeh.

A:6). On the ordered similarity matrix (Table 4:7) Tabiin Cl was found to be very similar to Skhiil 4. These results do not contradict those of Corruccini (1974a) or Stringer (1974b), but are different to those of earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986) where Tabiin Cl did not cluster with the "classic" Neanderthals from Europe or western Asia, but grouped with "pre or progressive" Neanderthal crania - Steinheim, Gibraltar 1 and Saccopastore 1 (see Condemi 1992; but also see Habgood 1985a).

The dating of the Tabiin Cl skeleton has another problem in that there are questions surrounding its stratigraphic position. Garrod and Bate (1937:64) recorded that Tabiin Cl was recovered from Layer C but "so near the surface of [Layer] C that the question must arise whether it does not represent a burial from level B" (but see Gargett 1989). Ruhlin ( 1998) has argued that Tabiin C 1 has a very derived occipitomastoid area compared to the other western Asian Neanderthals, which supports a later date and the possibility it was an intrusive burial from Layer B. The attribution of Tabiin Cl to Layer C could be supported by the amino acid racemisation dates discussed above. It has also been proposed that Tabiin Cl may have actually derived from Layer D due to dipping of the strata near where it was recovered (Ronen 1976, A.J. Jelinek in Discussion section in Ronen 1982). Ronen (1976) suggested that it was the reddish colour of the deposit in and around the burial of Tabiin Cl as opposed to the yellowish colour of Layer D, that led to the attribution of Tabiin Cl to Layer C. Ronen (1976) pointed

The current results are more consistent with the morphology of Tabiin Cl than the earlier analyses in that it displays many Neanderthal features . It is not, however, a "classic" Neanderthal for, as discussed above, it does not display the full suite of Neanderthal characteristics (Table 5:6) and in many cases it displays those that it does have to a lesser degree than most of the other members of this group. In this Tabiin Cl is similar to crania such as Krapina C, Saccopastore 1 and Gibraltar 1, a relationship that was highlighted in earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986; see also Condemi 1992). These differences may be due to Tabiin Cl being 87

50

0

100

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CMl P4

cc Ch DV3 02

Ml APl 01

Figure 4:7. Dendrogram from sum of squares cluster analysis with Tabon CI , Amud 1 and Shanidar 1.

A2

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t (6 .9) • LIA

2 TPH

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• LIC

St • • Sk5

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ov3• .ch

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PDA •• ZYB •• MDL AUB" NLH

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Figure 4:8. Correspondence analysis with Tabon C 1, Amud I and Shanidar 1.

88

chronologically earlier than most of the other western Asian and "classic" Neanderthals (but see Hublin [ 1998]. Problems of reconstruction may also have contributed to the differences). However, in general, Tabiin Cl and the other western Asian Neanderthals are more "generalised" or "less specialised" than their European counterparts.

to some western European Neanderthals such as those from Spy and Regourdou than to the Tabiin Cl mandible. Quam and Smith ( 1998) stressed the modern morphological components evident on the Tabiin C2 mandible, especially the presence of a chin, while also acknowledging the Neanderthal features. They proposed that Tabiin C2 has a strong phylogenetic connection with the Skhiil and Jebel Qafzeh hominids, but the morphology also reflects a contribution from Neanderthals. Dental metrics aligned Tabiin C2 with western Asian and European Neanderthals and separate it from Skhiil, Jebel Qafzeh and Upper Palaeolithic material.

Trinkaus (1983a, 1984a) suggested that Tabiin Cl was similar to other early Neanderthals from western Asia, especially Shanidar 2 and 4. As outlined earlier, there are also similarities between Tabiin Cl and Zuttiyeh in the supraorbital region and the degree of facial prognathism. There are also similarities in the supraglabellar region with Zuttiyeh, Skhul 4 and 9 and Jebel lrhoud I.

Stringer et al. (1979) placed Tabiin in their Homo sapiens Grade 3a with other Neanderthal material. Tabiin Cl is similar to other western Asia and European Neanderthals. Even though Tabiin C2 does display some features leaning towards later Neanderthals, it was still separated from them on the analysis of mandibular non-metric features (Fig. 5: 17). The Tabiin C2 mandible is also larger, more robust and morphologically different from the mandible of the Tabun Cl Neanderthal. Overall, Tabiin C2 is more aligned with archaic Middle Pleistocene hominids from Africa, Europe and Asia than with Neanderthals and can be regarded as an archaic hominid with affinities to both the Neanderthals and anatomically modem Homo sapiens.

On a bivariate plot of femoral head diameter (squared) against pubic length, Tabiin CI was positioned with Skhiil 9, Krapina 208, Kebara 2 and La Ferrassie l, and was separated from Skhiil 4 Jebel Qafzeh 9 and Upper Palaeolithic material (Rosenberg 1998:Figure 3). This confirms other observations that Tabiin Cl has Neanderthal features of the postcranial skeleton. On the analysis of mandibular non-metric traits (Fig. 5: 16) the Tabiin Cl mandible clustered with other Neanderthals including Shanidar l and La Ferrassie l. The Tabiin C2 mandible was also included in the analysis of non-metric traits. It was separated from the Tabiin C 1 mandible, and fell with Arago 2 between Neanderthal mandibles and the mandibles of early modern crania including that of Skhiil 5 (Fig. 5: 16). This position fits with the overall morphology of Tabiin C2 in that it displays Neanderthal and nonNeanderthal (more archaic) features.

Amud At present, up to sixteen individuals have been recovered from the Formation B within the Middle Palaeolithic levels at Amud Cave in the Wadi el'Amud near Lake Tiberias, Israel (Fig. 4:1; Hovers et al. 1995; Oakley et al. 1975; Suzuki and Takai 1970). The majority of the hominid material consists of isolated teeth and fragmentary postcranial material and there appears to be a predominance of infants and young children, although it has been suggested that this may be the result of systematic sieving during excavation (Hovers et al. 1995). It has been argued that three individuals preserve enough morphological detail to allow assessment of taxonomic attribution (Hovers et al. 1995). These are Amud 1(I), 2(II) and 7. The other fragmentary material from Amud will not be dealt with here, but details and descriptions of it may be found in Suzuki (1970b) and Hovers et al. (1995).

These placements of the Tabiin mandibles are similar to a metrical study by Roth (1983, 1985). A discriminant function analysis of metrical data classified Tabiin C2 as an "early modern" and not a Neanderthal (Quam and Smith 1998). Principal components analyses of dental metrics aligned Tabiin C2 with western Asian and European Neanderthals and separated it from Skhiil, Jebel Qafzeh and Upper Palaeolithic material (Stefan and Trinkaus 1998). A similar result was obtained through discriminant analyses of dental metrics (Stefan and Trinkaus 1998). The phylogenetic placement of the Tabiin C2 mandible has varied depending on the significance placed on particular morphological features. Trinkaus (1983a, 1995; Stefan and Trinkaus 1998) has argued that Tabiin C2 is best viewed as a late archaic hominid with affinities to the Neanderthals, because it has relatively large teeth, especially the anterior teeth, and displays Neanderthal features on the posterior segment of the mandible (mental foramen under first molar, H-0 mandibular foramen form, slight retromolar space). It is also argued that there is only slight development of a chin on the symphyseal region of Tabiin C2 that is closer to Amud 1 than Skhiil and Jebel Qafzeh (Stefan and Trinkaus 1998). Rak (1998), however, aligned Tabun C2 with anatomically modem Homo sapiens based predominantly on the presence of a chin and on his argument that Tabiin C2 only has a retromolar spaces because of marked pre-angular notches, which is different to the Neanderthal pattern. Rak (1998), aligned Tabun Cl with Neanderthals. Schwartz and Tattersall (2000) suggested that Tabiin C2 displayed more similarities

Amud I consists of an adult male skeleton (Suzuki 1970b). The cranium (Plates 4, 5 and 7), which was reconstructed using the Shanidar material to approximate missing portions, is large in overall dimensions (length, breadth and height) with a cranial capacity of 1740cc (Suzuki 1970b:Table VIIII) The cranium is long and narrow with a large moderately prognathic and heavily reconstructed face. The frontal is receding and gently curved, as are the parietals. In occipital view the cranial vault is rounded in profile. The teeth are large, especially the anterior teeth. Amud I displays many typical Neanderthal characteristics including a mastoid protuberance, occipital torus and a suprainiac fossa (Table 5:6; Santa Luca 1978; Suzuki 1970b). However, in some features Amud 1 differs from the typical Neanderthal pattern. It has a higher cranial vault, a more strongly curved sagittal contour, a less angulated occipital bone, little lambdoidal flattening or occipital bunning, a higher and more curved squamous margin of the temporal bone, larger mastoid 89

provided dates of 52,000+/-4,000 years BP (EU) and 66,000+/-5,000 years BP (LU). Preliminary TL dates of 5060,000 years BP for Formation B are reported by Rak et al. (1994) for Level B2/8 . Valladas et al. (1998) have provided a TL age range of 50-70,000 years BP obtained from burnt flints from levels B2 and B4.

processes , less midfacial prognathism, the orbits are not as rounded and the supraorbital torus displays a tendency to separate into superciliary and supraorbital elements . The mandible of Amud I (Plate 8) is large with big teeth, especially the anterior teeth and displays Neanderthal features (Table 5:6) including a retromolar gap and the mental foramen positioned under the first molar. The coronoid processes are large and higher than the condylar processes and the mandibular notch is relatively shallow with the deepest point near the condyle. The mandible, like the cranium, displays a number of non-typical Neanderthal features such as a more vertical ramus and a vertical symphysial region with slight symphyseal prominence or chin development (but see Schwartz and Tattersall 2000). Trinkaus ( 1983a; Stefan and Trinkaus 1998) stated that the Amud 1 mandible lacked the horizontal-oval mandibular foramen pattern, whereas Suzuki ( 1970b) described the left ramus as having this pattern and the right lacking it.

The fauna from Amud Formation B has been correlated with Layer F from Mugharet el-Wad , Mount Carmel , levels XIXXXV at Ksar 'Akil, and Shanidar Layer D (Suzuki 1970a; Suzuki and Takai 1970). At Amud there appears to be at least three intentional burials (see Hovers et al. 1995): Amud 1-an articulated skeleton in a flexed position with the head oriented towards the northwest ; Amud 7-an articulated skeleton of a young child with the head oriented towards the northwest and a possible associated grave good (red deer maxilla) ; and Amud 9-an articulated foot and lower leg. Hovers et al. (1995) have observed that the spatio-temporal distribution of the hominid remains at Amud is restricted in time and space in that the hominid remains are predominantly from level B 1 or B2 and are localised in distribution (Hovers et al. 1995:Fig. 1). This pattern is interpreted as "reflecting a kind of traditional , intentional burial/disposal ground" (Hovers et al. 1995:56).

The Amud I postcranial skeleton displays the Neanderthal configuration , although the crural index is higher and the estimated stature greater indicating that Amud 1 was a relatively tall individual (Table 4.8, Endo and Kimura 1970; Trinkau s 1981). Amud 2 is a fragment of the alveolar process of the right maxilla of an adult that closely resembles the morphology of Amud 1 (Hovers et al. 1995; Suzuki 1970b).

Gargett (1989, 1999, 2000) however , has argued that natural depositional circumstances can explain the completeness and state of articulation of these individuals and that there is no conclusive evidence for intentional burials at Amud. Whilst there are no clear burial pits at Amud , although Amud 7 was recovered from a small niche in the cave's north wall, Gargett's arguments do not conclusively demonstrate that natural burial was any more probably than intentional burial and so, as acknowledged by Gargett (2000), both options remain possibilities .

Amud 7 consists of the partial skeleton of a young infant possibly 10 months old (Rak et al. 1994). The facial skeleton is not preserved, but the mandible , portions of the temporal , parietal and occipital bones and the cranial base and postcranial elements are present. The mandible is relatively broad and anteriorly gently curved with large mental foramen. The mandibular corpus does not preserve a chin (Schwartz and Tattersall 2000). The foramen magnum is elongated and oval shaped and a well-preserved occipital torus is reported as being present (Hovers et al. 1995; Rak et al. 1994).

The stone artefacts from Formation Bat Amud were initially referred to as a transitional industry between Middle and Upper Palaeolithic industries (Suzuki 1970a; Watanabe 1968, 1970). However, A.J. Jelinek (1982b) has equated a limited number of artefacts from Amud with those from Layer D at Tabun and suggested that the artefacts were early in the technological chronology of the Mousterian in the Levant. Overall, however , the lithic industry at Amud is typical Mousterian produced by Levallois technique and is not a transitional industry, and is similar to Tabun B-type (Bar-Yosef 1992, 1998).

Early attempts at uranium-series and radiocarbon dating at the site did not provide acceptable results probably due to contamination (Grun and Stringer 1991; Suzuki and Takai 1970). Initial attempts at electron spin resonanc e (ESR) dating at Amud produced dates ranging from 65,000 to 106,000 years BP (Ikeya 1986), which are substantially older than more recent dates proposed for the site. The dating of the Middle Palaeolithic levels at Amud has been clarified with more recent thermoluminescence (TL) and ESR methods that have provided comparable results. Grun and Stringer ( 1991) reported ESR results on an artiodactyl tooth collected from near the top of Formation B of 42,000+/-3,000 years BP and 41,000+/-3,000 years BP for early uptake (EU) and 49,000+/-4 ,000 and 50,000+/-4,000 years BP for linear uptake (LU). Schwarcz and Rink (1998:Table 2) obtained ESR ages for Formation B ranging from 43,000 to 110,000 years BP (EU) and 48,000 to 113,000 years BP (LU). A tooth from level B 1/6, from which Amud 1 was recovered (Hovers et al. 1995:Table 1), provided dates of 43,000 +/-5,000 years BP (EU) and 48,000+/-6,000 years BP (LU). Level B2/8, from which Amud 2 and 7 were recovered (Hovers et al. 1995:Table 1),

Amud 1 was found to cluster, initially, with Shanidar 1 and La Ferrassie 1, and then with the other archaic hominid crania in the analyses (Fig. 4 :7, Table 4:4). On the correspondenc e analysis (Fig. 4:8), Amud 1 is again placed near Shanidar 1 and they are positioned in the same vicinity as the other Neanderthal crania . There were high loadings on the variables that document the sagittal contour of the posterior cranium and the size of the face (Fig. 4:8, Table A:6) . On the analyses of both cranial (Fig. 5:15) and mandibular (Fig. 5:16) non-metric traits Amud 1 was placed in an intermediate position between archaic and modem crania . On the mandibular analysis Amud 1 grouped with Ehringsdorf 6 (Fig. 5:16). In earlier analyses Amud 1 also grouped with Shanidar 1 and then the European Neanderthals 90

(Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). These results are in general agreement with those of Corruccini (1974a) and Stringer (1974b, 1978). In an analysis of the frontal bone, Simmons et al. ( 1991) found Amud 1 to be linked most closely with Zuttiyeh and, slightly further removed Tabiin Cl. Amud 1 was separated from Shanidar 1 and 5, unlike on the analyses discussed above. This separation in an analysis focusing on the frontal may be reflecting the strong possibility that Shanidar 1 and 5 may have experienced cranial deformation (Trinkaus 1982c, 1983a).

into consideration, the overall morphology of the Amud hominids fits within the Neanderthal pattern.

Mugharet el-Kebarab-Kebara The fragmentary remains of over 20 adult and sub-adult individuals have been recovered from the LevalloisoMousterian levels of Kebara cave, 13km south of the Mount Carmel caves, Israel (Fig. 4:1; Arensburg et al. 1985; BarYosef et al. 1986; Bar-Yosef et al. 1992; Rak and Arensburg 1987; Smith and Arensburg 1977). Only the two most complete and morphologically most diagnostic individuals will be discussed here.

The results from the multivariate analyses are consistent with the morphology of Amud 1, which displays Neanderthal features and is most similar to the western Asian Neanderthal Shanidar l. The results also highlight the morphological features that Amud 1 displays which are not typical of the Neanderthals. Suzuki ( 1970b) suggested similarities between Amud 1 and Skhul 4 in cranial dimensions, the form and development of the supraorbital region and in various features of the face, but it was noted that Skhul 4 was more advanced. Skhul 4 was also found to group with the Neanderthals on occasions (Fig. 4:3, 4:4), due to the sagittal contour of Skhul 4 and not its overall morphology. Thoma (1964, 1973) argued that Amud 1, along with other western Asian Neanderthals (Shanidar and Teshik-Tash), formed part of the ancestry of eastern Asian Mongoloids (see also Pichardo 1978). There is no morphological or metrical data that strongly supports this contention with respect to Amud 1. Stringer et al. ( 1979) placed Amud in their Homo sapiens Grade 3a with other Neanderthals.

From Unit X within the lower Mousterian levels the fragmentary remains of a 7-9 month old infant, Kebara 1, were recovered (Smith and Arensburg 1977). The remains consist of cranial fragments, portions of the maxillae and mandible and parts of the postcranial skeleton. Morphologically this individual was reported as resembling the immature individuals from Subalyuk, Gibraltar (Devil's Tower), Pech de l'Aze and Engis (Smith and Arensburg 1977), although more recent reports suggest that the remains are nondiagnostic along with the other fragmentary remains or isolated dental material (Bar-Yosef et al. 1992). Smith and Arensburg ( 1977) record that Kebara 1 was an intentional burial, but Gargett (1989,1999) has questioned this interpretation. More recent excavations at the site have produced a hominid skeleton from Unit XII within the Mousterian levels at the site (Arensburg 1989; Arensburg et al. 1985; Bar-Yosef et al. 1986; Bar-Yosef et al. 1992; Rak and Arensburg 1987). The remains consist of much of an adult male postcranial skeleton including the pelvis and the upper limbs, a complete mandible with teeth, the right upper third molar and the hyoid bone. The cranium and most of the lower limb bones were missing. The lower portion of the badly preserved left femur had been removed, unknowingly, during earlier excavations at the site. In general, the morphological features of the postcranial skeleton of this individual align it with the western Asian Neanderthals from Tabun C, Amud and Shanidar (Arensburg et al. 1985; Bar-Yosef et al. 1986; BarYosef et al. 1992; Rak 1990). The very robust mandible also displays Neanderthal features such as a retromolar gap, a receding symphysis, and the mental foramen positioned under the first molar. The Kebara 2 mandible, along with Tabun C2, has the horizontal-oval mandibular foramen form that is not common on western Asian hominids. Bar-Yosef et al. (1992) describe Kebara 2 as displaying a combination of plesiomorphic traits, especially the overall robustness of the individual, Neanderthal features on the mandible, upper limbs and pelvis, and morphologies such as on the hyoid, ribs and vertebrae that can fit into a modem range of variation.

Rak et al. ( 1994) have argued that an anteroposterior elongated and oval shaped foramen magnum is a Neanderthal autapomorphic trait as it was found on adult and immature Neanderthals, where preserved including Amud 7. CreedMiles et al. ( 1996) and Tiller ( 1998) have questioned the identification of this morphology as a Neanderthal autapomorphic feature because there is variation in the shape of the foramen magnum within modem samples. There also appears to be significant measurement variation when measuring the foramen magnum and determining the FM index (compare Creed-Miles et al. 1996:Table l; Tiller 1998:Table 3; and data in Rak et al. 1994). Even with these reservations, the morphology of Amud 7 suggests that it is the remains of an immature Neanderthal. Arensburg and Belfer-Cohen (1998; see also Arensburg 1989, 1991) have argued that Amud 1 differs from the Neanderthals in many morphological details including cranial height, supraorbital form, temporal bone curvature and large mastoids. However, a number of these differences, such as cranial height and frontal curvature, could be a result of the large cranial capacity of Amud 1 ( 1740cc see Suzuki 1970b:Table VIII-I) Amud 1 also has the long globular cranial shape of Neanderthals that is roundish in occipital view (Plates 4, 5, 7). The Amud 1 mandible has retromolar spaces and mental foramen under the first molar (Plate 8). A cladistic re-evaluation of the Amud 1 cranium and mandible found a number of Neanderthal apomorphies (Hovers et al. 1995).

The skeleton of Kebara 2 lay on its back in a shallow pit with the right arm across the chest and the left hand placed on the abdomen. The mandible rested on its base on the upper right shoulder and the cervical vertebrae were in place (Arensburg et al. 1985; Bar-Yosef et al. 1992). The excavators have concluded that Kebara 2 was an intentional burial and suggested that the skull may have been deliberately removed following the decay of the atlanto-occipital ligaments (BarYosef et al. 1992). Gargett (1999) has questioned these

There is morphological variation within the Neanderthals, especially those from western Asia and when this is taken 91

conclusions and argued that the burial pit was an erosional unconformity into which the body came to rest and where it was naturally covered by sediments, and that the cranium was displaced because it was exposed longer than the rest of the body.

It is worth noting that at Kebara there appears to be a clustering of hominid remains, especially of infants and children, in the proximity of the north wall of the cave suggesting a possible "dumping zone"(Bar-Yosef et al. 1992; Hovers et al. 1995:Fig. 1). The burials of Kebara I and 2 were positioned away from this cluster, with the former slightly south of the "dumping zone" and the latter placed more towards the centre of the cave (Hovers et al. 1995:Fig. 1). Hovers et al. (1995) have suggested that the place of disposal of the bodies (and one assumes method of treatment as well) may have shifted with time - burial in the centre of the cave (Unit XII) then near the north cave wall in (Units XI-IX) shifting to near the cave entrance in latter times (Units VNI).

The dating of the Kebara hominids has been established by use of a number of different dating techniques (Bar-Yosef 1998:Table 1). Radiocarbon dates of 35,300+/-500 years BP (GrN-2551) and 41,000+/-1000 years BP (GrN-2561) have been obtained for the upper Levalloiso-Mousterian level at the site (Vogel and Waterbolk 1963), but there could be problems with these dates due to the prolonged exposure in the section of the samples used (Bar-Yosef et al. 1986).

The affmities of the adult skeleton Kebara 2 are with Tabun C 1, Amud I and 7 and Shanidar 1 and 5 and contrast with the hominids from Skhul and Qafzeh (Bar-Yosef et al. 1986; Bar-Yosef et al. 1992; Hovers et al. 1995; Rak 1990). The morphology of the Kebara hominids fits within the Neanderthal pattern (but see Arensburg 1989, 1991 and Arensburg and Belfer-Cohen 1998).

Thermoluminescence (TL) average age estimates ranging from 48,300+/-3,500 years BP to 61,600+/-3,600 years BP have been obtained from 3 8 burnt flints from Mousterian units VI-XII (Valladas et al. 1987; Valladas et al. 1998). Unit XII that contained the adult hominid skeleton Kebara 2, was dated to 59,900+/-3,500 years BP , whereas Unit XI, from which the burial pit for Kebara 2 was cut, was dated to 60,000+/-3,500 years BP (Valladas et al. 1987). It is worth noting that Units XI and XII had younger average age estimates than Unit X, which was dated to 61,600+/-3,600 years BP (Valladas et al. 1987:Fig. 1). Isochron analysis of flints from Units XI and XII provided an age of 57,000+/8,000, which is close to the TL dates on the individual flints (Valladas et al. 1998).

Dederiyeh The remains of a two-year-old infant have been recovered from Dederiyeh Cave, northern Syria (Akazawa et al. 1995a; Akazawa et al. 1995b). The remains were from levels with lithic material similar to Tabun B-type Mousterian (Akazawa et al. 1995a; Akazawa et al. 1995b). The infant was lying on its back with arms extended, legs flexed and what may be "grave goods" in the form of a piece of flint positioned near the heart and a limestone slab near the head (Akazawa et al. 1995a; Akazawa et al. 1995b). The well-preserved skeleton consists of a very complete postcranial skeleton, mandible and cranial fragments. The individual displays Neanderthal features on the cranial and postcranial skeleton including a suprainiac fossa and transverse torus on the occipital, "swept-back" appearance of the zygomatic bone, receding symphysial profile on the mandible, shovelling of the upper central incisor and taurodontism of the first deciduous molar (Akazawa et al. 1995a; Akazawa et al. 1995b; Dodo et al. 1998). Many similarities have been identified with other Neanderthal infants, especially Pech de 1'Aze (Dodo et al. 1998).

Average electron spin resonance (ESR) ages on tooth enamel from Unit X at Kebara are reported as 60,000+/-6,000 years BP for early uptake (EU) and 64,000+/-6,000 years BP for linear uptake (LU) (Bar-Yosef 1998; Bar-Yosef et al. 1992; Griin and Stringer 1991), which are compatible with the TL average age estimate of 61,000+/-3,600 years BP obtained by Valladas et al. ( 1987). Porat et al. (1994) undertook ESR dating on five samples of burnt flint dated by Valladas et al. ( 1987). The average ages from the two radiation sensitive signals observed were: E' 48,500+/-5,100 years BP and Al - 64,600+/-12,000 years BP. The E' signal results were found to be consistently lower than the A I signal ages and the comparable TL dates. Units containing an early Upper Palaeolithic industry have provided radiocarbon dates of 46,000 years BP (Unit V) and 42,000 +/-l,800 years BP (Unit IV)(Bar-Yosef et al. 1992).

The excavators concluded that the remains were from an intentional burial (Akazawa et al. 1995a; Akazawa et al. 1995b). Gargett ( I 999) however, has argued that the Neanderthal remains were either the result of a rock fall or entered the cave through a "chimney" opening in the cave roof and came to rest in a "local low spot, depression, or pit" and were rapidly buried naturally ( a fall or washed in). There is no doubt that, as pointed out by Gargett ( 1999), the position of the skeleton is more suggestive of an accidental or unintentional placement of the body (possibly caused by some traumatic event such as crushing or a fall), than an intentional burial. However, intentional burial remains a distinct possibility for explaining the preservation of the skeleton of the two-year-old infant at Dederiyeh Cave.

The Kebara burial pit appears to have been cut down into Unit XII from lower part of Unit XI, where there were lateral hearths that overlay the burial and show no signs of truncation that would have resulted if the pit had been dug through them (Bar-Yosef et al.1992). The dating results support the contention of the excavators that the Middle Palaeolithic deposits at Kebara accumulated rapidly. An oxygen isotopic stage 3 age for the Kebara hominids is supported by the various dating methods used at the site. The lithic material from Kebara can be equated with that from the "chimney" deposits and Layer B at Tabun. (BarYosef et al. 1992; Jelinek, A.J. 1982a, 1982b). 92

•Kostenki

Kiik-Koba

~ ~ •

Dmanisi

Teshik. Tash.

•shanidar

Figure 4:9. Map showing western and central Asian sites mentioned in text.

It is generally stated that the Shanidar hominids were intentionally buried (Oakley et al. 1975; Solecki 1963; Trinkaus 1983a), however this assumption has been questioned on geomorphological and sedimentological grounds (Gargett 1989). The intentional burial scenario has included both individual burials (Shanidar 1, 2, 3, 5 and 7) and a single multiple burial (Shanidar 4, 6, 8 and 9), with "ritual activity" including piling rocks on top of a crushed individual (Shanidar 1) and the placement of flowers in a grave pit (Shanidar 4). Gargett (1989 and reply to Solecki 1989) has dismissed these conclusions and argued that the preservation of the individuals could be explained by episodes of ceiling collapse and rockfalls causing natural burial and entombment with wind blown pollen and flowers. Solecki ( 1989) has reaffirmed his earlier conclusions with respect to burials at Shanidar Cave, especially with respect to the Shanidar 4, the "flower burial".

Shanidar At the site of Shanidar Cave, northeastern Iraq, the remains of nine hominids have been recovered from Middle Palaeolithic levels (Fig. 4:9; Solecki 1957, 1961, 1963, 1971). The numbering of the individuals will follow that in the Catalogue of Fossil hominids: Part III (Oakley et al. 1975; Trinkaus 1983a). The hominids fall into two stratigraphical groups: Shanidar 2, 4, 6, 8, and 9 are stratigraphical lower than Shanidar 1, 3, and 5 and presumably older. Only the most complete crania will be dealt with here, but morphological descriptions of the other material, including the typical Neanderthal postcranial material (although it has higher brachia! indices, Table 4:8), are available elsewhere (Senyurek 1957a, 1957b; Solecki 1961, 1963, 1971; Stewart 1958, 1961a, 1963, 1977; Stringer and Trinkaus 1981; Trinkaus 1977a, 1977b, 1978a, 1978b, 1981, 1982b, 1983a, 1984a, 1995; Trinkaus and Stewart 1980; Trinkaus and Thompson 1986; Trinkaus and Zimmerman 1982; and references therein).

Shanidar 1 is an adult male skeleton (Solecki 1961; Stewart 1958; Stringer and Trinkaus 1981; Trinkaus 1983a, 1984a). The cranium is long and narrow and displays most of the typical Neanderthal features (Table 5:6). The cranial vault is higher than most Neanderthals, which is a result of the 93

the left side located under the 1st molar and two posteriorly placed small mental foramina on the right side.

elevation of its very arched parietals . The parietals are of modem proportions (Stringer and Trinkaus 1981). The frontal bone is very flat which, when coupled with the form of the parietals, suggests the possibility of cranial deformation (Trinkaus 1982c, 1983a; but see Brown 2003). The form of the occipital region on Shanidar 1 is typically Neanderthal, except it displays only slight development of an occipital bun (Santa Luca 1978; Trinkaus 1983a). The facial region also exhibits Neanderthal features, except it has minimally inflated maxillae and orbits that are not as rounded as those of other Neanderthals . Except for the lack of the horizontal-oval mandibular foramen pattern, the Shanidar 1 mandible displays the Neanderthal configuration (Table 5 :6). Shanidar 1 has anteroposterior elongated and oval shaped foramen magnum, which Rak et al. ( 1994) have argued is a Neanderthal autapommphic trait as it was found on adult and immature Neanderthals, where preserved (but see Creed-Miles et al. 1996; Tiller 1998).

Shanidar 5 consists of the fragmentary remains of an adult male (Stringer and Trinkaus 1981; Trinkaus 1977b, 1978b, 1983a, 1984a). The skull is predominantly represented by the anterior half of the cranium. The frontal bone is quite flat, while the parietals are markedly arched and the lambda is in a high position . The vault bones are very thick. Stringer and Trinkaus ( 1981) highlighted the unusual parietal curvature of Shanidar 5. Except for a lack of an anterior mastoid tubercle, the mastoid region of Shanidar 5 displays Neanderthal features (Table 5:6). The face of Shanidar 5, which displays an overall Neanderthal configuration (Table 5:6), is very large and prognathic . The nasal aperture dimensions are extremely large. The prominent, but relatively thin, supraorbital torus tapers laterally . Because of the unusual cranial configuration of Shanidar 5, Trinkaus ( 1982c, 1983a) suggested that like Shanidar 1, it had experienced cranial deformation . Problems with the reconstruction caused by extensive postmortem crushing also need to be considered when assessing and explaining the cranial configuration of Shanidar 5. Brown (2003) recorded that Trinkaus (pers com.) had advised that the high parietal curvature of Shanidar 5 was an artefact of the parietal being misaligned. Brown (2003) is of the view that whilst the frontal of Shanidar 5 is low, it does not display the degree of mid-frontal flattening or the presence of a pre-bregmatic eminence as evident on crania with cranial deformation .

Shanidar 2 is an adult male skeleton (Stewart 1961a; Trinkaus 1983a, 1984a). The skull is fragmentary and consists, primarily, of the right side of the cranium and the mandible. The occipital squama displays a typical Neanderthal pattern, with an occipital torus and suprainiac fossa. The mastoid region does not display all the typical Neanderthal features in that the mastoid process is quite large and is longer than the occipitomastoid crest, the mastoid crest is not continuous with the occipital torus and there is no anterior mastoid tubercle. Shanidar 2 also has an anteroposterior elongated and oval shaped foramen magnum (Trinkaus 1995). The fragmentary facial skeleton provides only limited information. The supraorbital torus is robust and appears not to taper laterally. The interorbital region is broad, as would have been the upper face. The orbits would have been large and subrectangular in shape. The nasal region appears to have projected anteriorly. The degree of midfacial prognathism, as demonstrated by the position of the right zygomatic root above, and not posterior to, the second molar and the relatively small retromalar spaces on the mandible, is not as great as on Shanidar 1 and most other Neanderthals. The robust and inflated right malar bone projects anteriorly and laterally. The palate is quite broad . The mandible of Shanidar 2 displays typical Neanderthal features including retromolar spaces and posteriorly positioned mental foramina, but it has a broad, vertical ramus, a more vertical symphysial region and lacks the horizontal-oval mandibular foramen pattern (Smith 1978; Trinkaus 1983a).

Although there are major similarities between the Shanidar hominids, such as the absence of the horizontal-oval mandibular foramen pattern , there appear to be two morphological groups that may reflect temporal differences. The earlier sample, Shanidar 2 and 4, have more robust faces, supraorbital tori that do not taper laterally and less midfacial prognathism, while the later group, Shanidar 1 and 5, are essentially typical Neanderthals in overall configuration . Trinkaus ( 1983a) argued that the material documented the change from an early Neanderthal morphology to a more typical Neanderthal pattern. This change is most evident in the facial region. The Shanidar hominid material came from the 8.5m thick Layer D, which contained a relatively homogeneous Mousterian industry with low frequencies of Levallois products possibly as a result of the small size of available raw material (Bar-Yosef 1992; Solecki 1963). Layer C contained a lithic assemblage with similarities to Aurignacian industries from the Levant (Trinkaus 1983a).

Shanidar 4 consists of an incomplete adult male skeleton (Stewart 1963; Trinkaus 1983a, 1984a). The fragmentary cranium displays typical Neanderthal features (Table 5:6), such as prominent mastoid and supramastoid crests and an occipital torus. The upper face appears to have been broad with only slight midfacial prognathism, while the robust supraorbital torus does not taper laterally (Trinkaus 1983a). The mandible of Shanidar 4, which displays significant damage and alveolar resorption , is large and robust and displays many Neanderthal morphological features (Trinkaus 1983a). As with the other Shanidar mandibles, Shanidar 4 lacks the horizontal-oval mandibular foramen pattern (Trinkaus 1983a). For the analysis of mandibular non-metric traits, the Shanidar 4 mandible was scored as displaying a large retromalar space, but Trinkaus (1983a) indicates that if rd the 3 third molar had been present the retromalar space would have been small. There is a large mental foramina on

Radiocarbon dates around 34,500 years BP have been obtained for the lower part of Layer C (Trinkaus 1983a:Table 1), suggesting that Layer D and the Shanidar Mousterian may have ended between 40-45,000 years BP. Radiocarbon dates of 46,900 +/-l ,500 years BP (GrN-2527) and 50,600+/3,000 years BP (GrN-1495) have been obtained from the upper part of Layer D (Vogel and Waterbolk 1963), which provide an approximate date for Shanidar 1, 3 and 5. These dates also provide a minimum age for the other hominid material, Shanidar 2, 4, 6, 8, and 9, but this deeper material could be significantly older depending on the geomorphological history of Layer D.

94

Solecki (1963) suggested an age of 60,000 to 70,000 years BP for the middle of Layer D, and 100,000 years BP for the bottom. These dates were based on a climatic reconstruction for the site that was, through the use of radiocarbon detenninations from the upper part of the sequence, correlated to sequences from other sites in western Asia and by assuming a constant rate of deposition within the cave (Solecki 1963), although there is no compelling reason to assume a constant rate of deposition at the site. The Catalogue of Fossil Hominids (Oakley et al. 1975) records inferred ages of 46-47,000 year BP for Shanidar 1, 3 and 5 and 60,000 years BP for Shanidar 2, 4 and 6 (Shanidar 7 was noted as being stratigraphically the oldest of the hominids). Bar-Yosef ( 1992) suggested a tentative estimate that the Shanidar hominids could be placed in the period 50-70,000 years BP. What is clear is that there is a stratigraphic and therefore a temporal distinction between the two hominid samples, although the extent of this chronological separation is unknown at present (Fig. 4:2).

Simmons et al. ( 1991) undertook a multivariate analysis of the frontal bone of hominids from western Asia. On the plots from the principal components analyses of log size and shape and log shape Shanidar 1 and 5 were positioned nearest each other and well separated from the other western Asia hominids (Simmons et al. 1991 :Figures 1 and 2). On the UPGMA cluster analysis of log shape Shanidar 1 and 5 were late joiners to the other cluster of archaic hominids (Simmons et al. 1991:Figure 4). The separation of Shanidar 1 and 5 in analyses focusing on the frontal squama supports the contention that these individuals may have experienced cranial deformation (Trinkaus 1982c, 1983a). Trinkaus (1982a, 1983a) suggested that Zuttiyeh displayed morphological similarities to the fragmentary Shanidar 2 and 4 individuals. As noted earlier, Shanidar 2 and Zuttiyeh preserve mostly different regions of the cranium and so comparisons between them are difficult. Where comparisons can be made there do not seem to be any marked similarities. Shanidar 4, like Zuttiyeh, has a robust supraorbital torus that appears not to have tapered laterally, and a steep supraglabellar region, although the remainder of the frontal of Shanidar 4 is much flatter than that of Zuttiyeh.

Only Shanidar 1 and 5 were included in cranial analyses ( see earlier discussion of possible cranial deformation). Shanidar 1 initially clustered with Amud 1 and La Ferrassie 1, and then with the other archaic crania in the analyses (Figs. 4:7, 4:8, Table 4:4). On the correspondence analysis the Neanderthals are separated from the other archaic hominids, with Shanidar 1 and Amud 1 falling near each other (Fig. 4:8). There were strong loadings on the variables that document the sagittal curvature of the parietal and occipital bones and the size of the face (Fig. 4:8, Table A:6).

The placements of Shanidar 1 and 5 are consistent with the morphologies of the two individuals, which display most of the typical Neanderthal features. The analyses also highlight a close morphological similarity between the western Asian Neanderthals, especially Amud 1 and Shanidar 1 and 5. Thoma (1964, 1973) argued that the Shanidar material, along with other western Asian Neanderthals, (Amud and TeshikTash) was part of the ancestry of the eastern Asian Mongoloids (see also Pichardo 1978). There is no morphological or metrical data that strongly supports this contention.

On the analysis of mandibular non-metric traits (Fig. 5:16) the Shanidar mandibles were placed with the other Neanderthal mandibles, including those of Tabun Cl and La Ferrassie 1. Shanidar 5 grouped with other Neanderthal crania (Figs. 4: 10, 4: 11, Table 4:5). There were high loadings on the variables that dealt with the sagittal contour of the frontal and parietals and facial size (Fig. 4:11, Table A:7).

Suzuki ( 1970b) identified strong similarities between the Shanidar hominids and Amud 1, but also identified differences including the form of the supraorbital torus and maxillae, frontal and parietal curvature and less development of a chin on the mandible. Suzuki (1970b) was of the view that the Shanidar material resembled the classic Neanderthals more than the Amud material.

Table 4:5. K-means cluster analysis with Shanidar 5. Analysis includes 74.1% of the total within-group variance. Group 1 Combe Capelle, Chancelade, Dolni Vestonice 3, Oberkassel 1, Oberkassel 2.

As mentioned earlier, Trinkaus ( 1983a, 1984a) suggested similarities between the early Shanidar material and the other early western Asian hominids, Zuttiyeh and Tabiin Cl and C2, and classified them all as early Neanderthals. Because of the fragmentary nature of much of this material it is difficult to assess, adequately, this assumption, but the fossils do display at least some similarities in morphology, especially in the supraorbital region. However, as discussed previously, it is hard to justify an early Neanderthal classification for Zuttiyeh at least and Tabun Cl could be significantly younger than the early Shanidar material.

Group 2 Predmost 3, Predmost 4, Cro-Magnon 1. Group3 Steinheim, Kabwe 1, Skhiil 5. Group 4 Mladec 1, Abri Pataud 1.

Stringer et al. (1979) placed Shanidar into their Homo sapiens Grade 3a with other Neanderthal remains. Overall the Shanidar material displays most similarities to Neanderthals from western Asia and Europe.

Group 5 Gibraltar 1, La Chapelle 1, La Ferrassie 1, Shanidar 5. These results are comparable to those of Corruccini (1974a), Howells (1970) and Stringer (1974b).

95

50

0

100

St

Kl

Sk5 Gl LCl LFl

Sh5 P3 P4

CMl Ml

APl

cc Ch DV3 02 01

Figure 4:10. Dendrogram from sum of squares cluster analysis with Shanidar 5.

A2 (5.9) I

•rPH

2

Kl.

.S1160,000 years BP (U-Pa) for Arago 21, whereas Arago 47 provided ages of 357,000+nd-140,000 years BP (U-Th) and >135,000 years BP (U-Pa). Laitman (1985) reports that differences in dating results for Arago 21 and Arago 47 may be due to the latter being recovered from an area affected by external ground water.

Many attempts have been made to date the Arago deposits. Cook et al. (1982) provide a detailed discussion of the dating methods used and the results obtained at the site. The fauna is said to support both a "Riss" and a "Mindel'' age. There are problems with the use of the uranium series method at the site, but when used on carbonate samples and bone from the upper unit, "ensemble superieur IV", the method indicated that the unit was older than 350,000 years BP (Cook et al. 1982). Aspartic acid racemisation dates (using uranium series dates for calibration), on bone from the same deposit as Arago 21, provided an age range between 320,000 and 360,000 years BP. But as Cook et al. state ...the application of AAR dating to this site must await the resolution of the U-S problems. (1982:36)

As has been mentioned, there are problems with the use of the various methods at the site, which means that the numerous dates are questionable (Cook et al. 1982; Debenham 1983; Skinner 1983; Stringer et al. 1984). Cook et al. state The complicated nature of the cave site has caused problems for all dating techniques. (1982:37) A mid-Middle Pleistocene age, possibly 350,000 to 450,000 years BP, for the Arago hominids would seem a reasonable estimate . On the mandibular non-metric analysis (Fig. 5: 16), the two Arago mandibles were separated from each other. Arago 2 grouped with the Tabu.n C2 mandible, intermediate between the Neanderthal and anatomically modem mandibles. Closest to it were the Saint-Cesaire, Atapuerca 1, Guattari-Monte Circeo 2, and Skhul 4 mandibles. The Arago 13 mandible fell within a group of archaic mandibles including Baiiolas, Krapina G, and Guattari-Monte Circeo (Fig. 5: 16). This group of mandibles is intermediate between Neanderthal mandibles and a group of archaic mandibles made up of Mauer, Montmaurin, Malamaud 1, Hortus 4, and La Naulette

De Lumley et al. (1984) discuss the dates obtained from numerous dating techniques used at Arago. "Ensemble superieur IV" has produced a uranium series date of greater than 350,000 years BP, an amino acid date of 400,000 years BP, electron spin resonance dates of 500,000 and 240,000 years BP, and thermoluminescence dates of 450,000 and 230,000 years BP. "Ensemble superieur III" has provided a uranium series date of greater than 350,000 years BP, an electron spin resonance date of 450,000 years BP, and an amino acid date of 440,000 years BP. The lower travertine floor at the site has produced a uranium series date of greater than 350,000 years BP, electron spin resonance dates of 700,000 and 260,000 years BP, and thermoluminescence dates of 360,000 and 220,000 years BP.

1.

These groupings are consistent with the morphologies of the two mandibles. Due to its slight retromolar gap and mental foramen positioned under the first molar, Arago 2 can be regarded as being Neanderthal-like, which is consistent with its position on the plot from the correspondence analysis (Fig. 5:16). The tooth size of Arago 2 is also well within the 125

Kabwe 1 and possibly Bodo 1. In overall shape Arago 47 recalls the Swanscombe parietals .

Neanderthal range (Wolpoff 1980a, 1982). The broad and moderately high ascending ramus of Arago 2 is similar to that on the Tabiin C2, Mauer and Bafiolas mandibles. Apart from having a mental foramen under the first molar, Arago 13 does not display the typical Neanderthal morphological configuration (Table 5 :6). In the lack of a retromolar gap, receding symphyseal region without development of mental eminence, and overall size and robustness, Arago 13 recalls the Mauer jaw . It differs from Mauer in having a higher, narrower, and more posteriorly angled ascending ramus . Arago 13 is also similar to the Montmaurin mandible in the form of the ascending ramus, receding symphyseal region, and lack of a retromolar space. In the latter two features it is also similar to the Bafiolas mandible. Krapina G and Guattari-Monte Circeo 3 have a more developed retromolar gap than Arago 13. Similar relationships for these mandibles were discerned by Aguirre and de Lumley (1977) .

M-A. de Lumley (1975) regarded the Arago material as anteNeanderthals . Stringer et al. (1979) placed Arago with Steinheim, Swanscombe and Montmaurin in their Homo sapiens Grade 1 or 2. Dean et al. ( 1998) placed Arago within an early- pre-Neanderthal evolutionary stage with Petralona and Mauer . The Arago material is best regarded as archaic Homo sapiens of the mid-Middle Pleistocene.

Bilzingsleben Faunal material, pebble tools and hominid remains have been recovered from the Steinrinne travertine quarry near the village of Bilzingsleben Germany (Fig. 5:3; Mania 1976; Mania and Vlcek 1981). The cranial fragments and molar (Mania and Vlcek 1981; Stringer 1980, 1981; Stringer et al. 1984; Vlcek 1978a; Wolpoff 1980a, 1980b) were widely dispersed over the site and so may have been scattered by water. They may belong to a single individual as there are no duplication of parts or morphological inconsistencies.

As mentioned above, there is a certain degree of postmortem deformation of Arago 21, which influences some of the measurements (Plate 9). This was taken into consideration when the measurement s were taken. For example, only the relatively undeformed left orbit was measured . All measurements were also checked on a cast that had (to a large extent) been corrected.

Two occipital fragments (Al and A2) join to form a relatively complete bone that is very thick and strongly flexed. The occiput is broad and quite low, while the nuchal plane is flat. There is a large mound-shaped transverse torus that is most prominent medially. No suprainiac fossa is present, although there is a shallow supratoral sulcus that is not continuous at the mid-line. The inion is separated from the endinion , but coincides with opisthocranion .

Arago 21 grouped with Petralona and Kabwe 1, and then with the other archaic hominids . There were high loadings on the variables that document the form of the frontal and the size of the face. In other analyses, Arago 21 also grouped with Bodo 1. When a composite reconstruction (Plate 9, Arago 21, Arago 4 7 mirrored to form both parietals , the temporal of Sangiran 17, and the occipital of Swanscombe) of the Arago cranium (Pretirage , 1er Congres International de Paleontologie Humaine, Nice 1982, Vol. 1) was used, it also grouped with Petralona , Kabwe 1 and Skhiil 5. In earlier analyses Arago 21 was a member of a group that included Steinheim , Saccopastore 1, Gibraltar 1, Tabiin Cl, and Skhiil 5 (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). Stringer (1978) found that, although larger, the Petralona face was quite similar to that of Arago 21. Saccopastore 1 was also found to be relatively similar to Arago 1, whereas Kabwe 1 and Steinheim were not. Van Vark (1983) also found Arago 21 to be similar to Petralona, Kabwe 1, and Steinheim.

The glabellar fragment (B 1) is very robust. It indicates a very prominent supraorbital torus continuous across the glabella, and a broad interorbital area with a huge nasal root. The frontonasal suture forms an inverted "V". There is said to be no supraorbital sulcus or supraglabellar depression (Stringer 1981; Vlcek 1978a), although Mania and Vlcek (1981:141) stated that "there is a broad, smooth, and shallow depression" above the glabella "which blends with the surface of the frontal" , and Wolpoff (1980b : 348) stated that "A broad sulcus separates the supraorbitals from the frontal squama". The small segment of the frontal squama is very thick, which is consistent with the other frontal fragment (B2). The frontal sinuses are well-developed . The molar has a relatively large pulp cavity .

The results of the analyses are in general agreement with the morphology of Arago 21. In overall size, robustness , marked postorbital constriction, relatively flat frontal, form of the supraorbital torus, amount of facial prognathism (after correction) , wide interorbital region, pneumatisation of the malars , lack of canine fossae, and shelving of the zygomatics into the maxillae, Arago 21 does indeed resemble Petralona, Kabwe and Bodo 1 (Plate 1). The facial breadths of Arago 21 are similar to those of Petralona , although it is shorter. In facial height Arago 21 is similar to Steinheim.

There is some debate over the dating of Bilzingsleben. Mania and Vlcek ( 1981) stated that ...the travertine complex including the fossil bed can be placed in the Elster/MindelSaale/Riss interglacial, that is, the Holsteinian of north European glacial chronology . (1981:134) This is based on the geomorphology of the site, the fossil fauna and flora from the travertine , and on an amino acid racemisation date of 230,000 years BP (Mania and Vlcek 1981). Harmon et al. (1980) have obtained uranium series dates , for a calcite lens within the fossiliferous lower sandy travertine horizon, which provide an age of 228,000+ 17,00012,000 years BP for the hominid remains . They place the site in the penultimate interglacial, and equate it with oxygen isotop e stage 7, which dates between 190,000 and 245,000 years BP (Table 5:1; Griin et al. 1997). Svoboda (1987) report s additional uranium series dates of 179,000-301,000

Wolpoff (1980a, 1980b) suggests that, in overall facial size, Arago 21 is intermediate between Petralona and Steinheim . He also contends that the Arago face resembles that of Saccopastore 2. Bodo 1 has a similarly flat mid-facial region, whereas Petralona and especially Kabwe 1 have more midfacial projection (Plate 1). The form of the frontonasal suture of Arago 21 differs from that of Petralona, Bilzingsleben, and probab ly Steinheim, and is more similar to that of 126

years BP and 335,000-350,000 years BP and amino acid dates that range between 280,000 and 350,000 years BP.

classified as Homo erectus, but the remainder of the cranium demonstrates that it is an archaic Homo sapiens.

An alternative correlation of the site, based on floral and faunal material, with the warmer oxygen isotope stages 9 or 11 has been suggested (Cook et al. 1982). These stages date to between 295,000 and 333,000 years BP, and 375,000 and 415,000 years BP respectively (Table 5:1; Griin et al. 1997). Additional uranium series determinations have yielded ages of greater than 350,000 years BP (Cook et al. 1982; Schwarcz 1982). Schwarcz et al. (1988) have obtained uranium-series and electron spin resonance (ESR) age estimates that indicate that the age for the hominid-bearing deposits are less than or equal to 414,000 +/- 45,000 years BP and no later than 280,000 years BP. Whilst this would relate to oxygen isotope stage 11, they caution that oxygen isotope stage 9 cannot be excluded as a possible date (Schwarcz et al. 1988).

If the Bilzingsleben material is late Middle Pleistocene in age it is younger than Vertesszollos 2. This would present a marked morphological dichotomy in that the younger material presents the more archaic morphology. There are definitely differences between the Bilzingsleben material and other European archaic Homo sapiens, and similarities between it and Homo erectus material, but as Stringer states ...there is certainly equal justification for grouping the Bilzingsleben fossils with European middle Pleistocene specimens ... (1981:9) Stringer et al. ( 1979) placed the Bilzingsleben remains in their Homo sapiens Grade 1, along with Petralona, V ertesszollos, Mauer, Kabwe 1 and Bodo 1. Dean et al. (1998) placed Bilzingsleben within a pre-Neanderthal evolutionary stage with V ertesszollos, Atapuerca SH site, Swanscombe, Steinheim and Reilingen, which display incipient Neanderthal morphologies.

Again, a late Middle Pleistocene age for the Bilzingsleben hominid remains would seem probable, although an earlier date cannot be ruled out.

The Bilzingsleben remains are too fragmentary to allow an "exact" taxonomic placement of them. Overall, however, the hominid( s) can be regarded as a very robust male of morphologically variable population of the latter half of the Middle Pleistocene from Europe (Habgood l 989e ).

The Bilzingsleben hominid remains were too fragmentary to include in any analyses. The occipital bone is said to show greatest similarity to Sinanthropus 3 (the Locus E adolescent), Sangiran 17, and Olduvai H9, all Homo erectus crania (Mania and Vlcek 1981; Vlcek 1978a). Little similarity was seen between the Bilzingsleben occipital and those of Kabwe 1, Swanscombe, Steinheim, Ehringsdorf 9, or the Ngandong hominids, although some parallels were found on V ertesszollos 2 (Mania and Vlcek 1981; Vlcek 1978a). The Bilzingsleben occipital was not initially compared to Petralona (Mania and Vlcek 1981; Vlcek 1978a). This has been done, with Stringer (1980, 1981) suggesting that the occipital torus ofBilzingsleben was like a smaller version of that on Petralona. Wolpoff (1980a) also suggests that the Bilzingsleben occipital is similar to Petralona and V ertesszollos 2. Smith ( 1982) suggests that the F ontechevade calotte would be a good fit for the Bilzingsleben occipital.

Montmaurin A relatively complete mandible with six molars in place was recovered from a vertical fissure (la Niche) in the Miro cave system at Montmaurin, southwestern France (Fig. 5: 1; Billy and Vallois 1977; Cook et al. 1982; Oakley et al. 1971; Stringer et al. 1984). Other skeletal remains were recovered from Montmaurin, Coupe-Gorge (Oakley et al. 1971). The mandible (Billy and Vallois 1977; Stringer et al. 1984; Vallois 1956) is from a young adult female(?). It is very robust, while the teeth are relatively small, except the large third molars. The symphysial region is receding, the alveolar plane is well-developed and the ascending rami are relatively tall and straight with deep sigmoid notches and large coronoid processes. The large condyles are oval in shape (from above) and are not flattened. There are multiple foramina positioned below the fourth premolar-first molar. A retromolar gap is not present. The gonial angles are evenly curved. The mylohyoid groove is partly closed. The alveolar arcade is long, with only moderate posterior divergence and some anterior flattening. The molars are moderately taurodont.

The glabellar fragment is said to be most similar to those of Olduvai H9 and Sangiran 17, and quite different to those of Arago 21, Steinheim, and Ehringsdorf 9 (Mania and Vlcek 1981; Vlcek 1978a). Stringer (1980, 1981), however, argues that the glabellar morphology of Bilzingsleben is more similar to that of Arago 21 than had been suggested, and that it also resembles Petralona in mediosagittal section. Arago 21, however, has a nasofrontal suture that is more horizontal than that of Bilzingsleben. In this feature Bilzingsleben is similar to Petralona which also has a frontonasal suture that forms an inverted "V".

It is difficult to establish a date for the Montmaurin mandible because quarrying has removed much of the fill from "la Niche" and because it is difficult to equate the "la Niche" deposits with the caves in the Miro complex (Billy and Vallois 1977; Cook et al. 1982; Stringer et al. 1984). "La Niche" was probably filled by material from the grotte de la Terrasse, which is directly above it (Billy and Vallois 1977; Cook et al. 1982). Fauna and flora from the deposits suggest a penultimate interglacial age (Billy and Vallois 1977; Cook et al. 1982; Oakley et al. 1971; Stringer et al. 1984), although earlier dates have been suggested (Cook et al. 1982). Vallois ( 1956) suggested that a last interglacial age for the deposits was also possible. This has been supported by studies that

Mania and Vlcek (1981), Vlcek (1978a), and Vandermeersch ( 1985) regard Bilzingsleben as a Homo erectus, whereas Stringer ( 1980, 1981, 1985) and Wolpoff ( 1980a, 1980b) think it is an archaic Homo sapiens. Wolpoff (1980a, 1980b) suggests that European Middle Pleistocene males like Petralona and Bilzingsleben were very robust and resembled Homo erectus crania more than the females such as Steinheim and Swanscombe. Stringer ( 1980) also suggests that, if only the glabellar portion of the frontal and the occipital squama of Petralona were preserved, it would be 127

indicate a late "Riss" or early last interglacial age (Stringer et al. 1984).

reconstruction of the whole cranium . However, Wolpoff (1980a) is of the view that the cranium is extensively warped . He suggests that the whole left side is twisted toward the mid-line, and that the back of the cranium is warped underneath and to the side. Wolpoff ( 1980a:228) therefore , stated that "the skull must be analyzed with great caution" . From the use of computerised tomography ( CT) scans it has been suggested that the distortions are limited to the transition between the splanchnocranium and the neurocranium, and that they can be allowed for in reconstructions (Czarnetzki 1987; Laitman 1985).

Vallois (1956) stated that a pre-Mousterian type industry was associated with the mandible, whereas Boule and Vallois (1957) reported that no implements accompanied the mandible, but an archaic Mousterian industry was present in nearby deposits. Oakley et al. ( 1971) contend that the associated lithic material consisted of atypical flint and quartzite flakes. According to Billy and Vallois ( 1977), archaeological studies of the site support faunal and floral data that indicates a penultimate interglacial age.

The relatively small cranium is thought to be that of a young adult female. The gracile vault is long, low, moderately flattened , and narrow, with marked postorbital constriction , and only moderately thick vault bone s. The maximum cranial breadth is located near the inferior margin of the fairly vertical and flat parietals . The mastoid processes are small, the supramastoid crests are slight , and there are no occipitomastoid crests. The occipital is well-rounded , although Wolpoff (1980a) stated that this has been influenced by warping . There is little lambdoidal flattening , although there is a slight pre-lambdoidal depression . The transverse occipital torus is not excessively large. It is tripartite in structure with more prominent lateral segments and a wide, flattened medial portion above which there is a suprainiac fossa (Santa Luca 1978). There is no external occipital protuberance, the opisthocranion is located above the inion, and there is a lack of separation between the inion and endinion. The basicranium is well flexed (Laitman et al. 1979).

At present a conclusive date for the Montmaurin mandible is not available, but a penultimat e interglacial age would seem most parsimonious (Cook et al. 1982; Stringer et al. 1984). The Montmawin mandible was used in the analysis of nonmetric mandibular traits (Fig. 5: 16). It grouped with Mauer , Malamaud 1 and Hortus 4, which also have multiple mental foramina positioned anterior to the first molar . This group was well separated from both Neanderthal and anatomically modem mandibles (Fig. 5:16). The position of the Montrnaurin mandible is consistent with its overall morphology as it does display many similarities with these other mandibles, especially Mauer (Billy and Vallois 1977; Stringer et al. 1984; Vallois 1956). Montmaurin has narrower ascending rami and deeper sigmoid notches than Mauer. Montmaurin also resembles the Arago mandibles in some features (Billy and Vallois 1977; de Lumley, M-A. 1975). The molars are described as primitive, resembling those of Homo erectus (Billy and Vallois 1977; Vallois 1956). Billy and Vallois (1977) document other features (many of which relate to robustness) of Montrnawin that resemble Homo erectus mandibles. Although , Montmaurin does not have the characteristic Neanderthal retromolar gap or mental foramina positioned under the first molars, it does resemble the Neanderthals in a number of features (Stringer et al. 1984; Vallois 1956). Vallois (1956) argued that the overall morphology of Montmaurin was more similar to the Neanderthals than to Mauer, and that it had a Neanderthaloid morphology. He referred to it as preneanderthal (Vallois 1956). Stringer et al. ( 1979) placed Montmaurin with Steinheim, Swanscombe and Arago in their Homo sapiens Grade 1 or 2.

The relatively small face is delicate and not excessively prognathic, even allowing for the missing portion of the maxillae. The face is relatively flat, and does not display marked mid-facial prognathism. The supraorbital region is very pronounced and projects markedly forward. It forms a double arch that follows the contour of the orbits and thins laterally. A broad ophryonic groove separates it from the frontal squama, which has a relatively steep supraglabellar segment. The frontal sinuses are quite extensive . The right orbit is small and semi-rectangular in shape. The interorbital area is wide, and the nasofrontal suture appears to form an inverted "V". The nasion is depressed, and the saddle-shaped nasal bones are prominent. The nasal aperture is short and very broad. The gracile right malar has a shallow canine fossa (Boule and Vallois 1957; de Lumley 1976; contra Brace 1964). There is only slight angulation as the right malar shelves into the right maxilla. The remaining teeth are relatively small and display a moderate degree of taurodontism .

Steinheim A hominid cranium was found in the Sigrist gravel quarry at Steinheim, north of Stuttgart, West Germany (Fig. 5:3; Adam 1985; Howell 1960; Oakley et al. 1971). The cranium (Adam 1985; Howell 1960; Stringer 1980, 1981; Stringer et al. 1984; Wolpoff 1980a, 1980b) is almost complete , except for damage to the frontosphenoidal region on the left side, and missing parts of the left side of the face, the anterior segment of the maxillae, and part of the cranial base. It has been suggested that the damage to the left side of the vault was caused by "a blunt instrument" during life, and that the damage to the cranial base was the result of intentional enlargement of the foramen magnum for the removal of the brain (Adam 1985). Howell ( 1960) felt that the right side of the Steinheim cranium was perfectly preserved and permitted the accurate

W olpoff ( 1980a, 1980b) suggests that Steinheim was a female in a Middle Pleistocene population that had Petralona as a male . Stringer (1980, 1981), however , argues that the differences between these two crania are greater than one would expect if they were solely due to sexual dimorphism , and he suggests that there were evolutionary differences between them that are probably caused by temporal separation . The geology and fauna from the sandy loam unit, in which the skull was found, are generally thou ght to be Holsteinian in age (Adam 1985; Cook et al. 1982). Adam (1985:274) stated that Steinheim can be "securely correlated to the 128

Holstein or Mindel-Riss interglacial". However, there are some methodological problems with this attribution (Cook et al. 1982). It has been suggested that the site where the cranium was found need not be of the same age as that where the fauna was recovered (Butzer and Isaac 1975). This suggestion may be supported by the fluorine, uranium and nitrogen analyses (Oakley 1980; Oakley et al. 1971). Ikeya ( 1986) has obtained an electron spin resonance age range of 94,000(?) to 161,000(?) years BP for a bone sample from the Steinheim site. The Steinheim fauna is generally comparable to that from Swanscombe (Cook et al. 1982; Sutcliffe 1964). A late Middle Pleistocene age, somewhere around 300,000 years BP, for the cranium would seem most appropriate at present.

1984, 1985) found Steinheim to be close to Petralona, Kabwe 1 and Arago 21. The results are also consistent with the overall morphological configuration of the cranium in that it displays similarities to other mid- to late Middle Pleistocene crania ( see discussions of Petralona, Bilzingsleben and Swanscombe ). Steinheim, Petralona and Kabwe 1 also have well flexed cranial bases. Although they were not included in the same analysis, Steinheim displays many similarities to Swanscombe (Breitinger 1964; Corruccini 1974a; Howell 1960; Morant 1938; Stringer 1974b, 1978; Weidenreich 1943b; Weiner and Campbell 1964; Wolpoff 1980a, 1980b). The sagittal contour of Steinheim resembles that of F ontechevade 5. There are also some contrasts between Steinheim and the robust Petralona and Bilzingsleben remains. As discussed before, these may be due to a great degree of sexual dimorphism or chronological separation. In general, Steinheim displays a mixture of Neanderthal and nonNeanderthal features. Stringer states ...for me, at least, this specimen [Steinheim] sits less easily in the early Neanderthal group, since it apparently departs in a number of plesiomorphous retentions from the typical Neanderthal condition ... ( 1985:293-4)

Measurements for Steinheim were obtained from both corrected and uncorrected casts held in the Natural History Museum, London, so as to compensate for the warping of the cranium. Steinheim was one of the core group of crania that was used in most of the analyses (Table 1:2). It generally grouped with archaic Homo sapiens crania such as Petralona, Kabwe 1, Skhul 5, Jebel Qafzeh 6, Jebel Irhoud 1, Dali and Maha (Figs. 5:7, 5:8, 5:9, 5: 10, 5: 11, 5: 12, 5: 13, Tables 5:3, 5:4, 5:5). There were high loadings on the variables that document cranial curvature, and facial size (Figs. 5:8, 5:11, 5:13, Tables A:10, A:11, A:12). On occasions Steinheim clustered with Saccopastore 1 (Fig. 2:5, 2:6, 6:2), Gibraltar 1 (Fig. 4:8), or with a group of Neanderthal crania (Fig. 4:9). On the analysis of non-metric cranial traits Steinheim fell on its own (Fig. 5:15). Positioned closest to it were Kabwe 1 and Spy 1 (Fig. 5: 10). Similar groupings were found in earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986).

Steinheim has a non-Neanderthal face in that it has a subrectangular orbit, a canine fossa, and only limited mid-facial prognathism, but it does have a tripartite occipital torus with a suprainiac fossa that is typical of Neanderthals (Table 5:6; Hublin 1984; Santa Luca 1978; Stringer 1978). Both Howell (1951) and Vallois {1954, 1958; Boule and Vallois 1957) regarded Steinheim as a "progressive" or "pre-Neanderthal". Stringer et al. ( 1979) placed Steinheim with Swanscombe, Arago and Montmaurin in their Homo sapiens Grade 1 or 2. Dean et al. (1998) placed Steinheim within a pre-Neanderthal evolutionary stage with Bilzingsleben, V ertesszollos, Atapuerca SH site, Swanscombe, and Reilingen, which display incipient Neanderthal morphologies.

Table 5:4. K-means cluster analysis of European core group. Analysis includes 86.6% of the total within-group variance. Group 1 Steinheim, Kabwe 1, Gibraltar 1, La Chapelle 1, La F errassie 1, Skhul 5.

Swanscombe Group 2 Combe Capelle, Chancelade, Dolni Vestonice 3, Oberkassel 1, Oberkassel 2.

Two parietals and an occipital squama, along with Acheulean artefacts and faunal remains, were recovered from Bamfield gravel pit near Swanscombe, England (Fig. 5: 1; Oakley et al. 1971; Ovey 1964). The three bones articulate to form a relatively complete posterior cranium of what is generally thought to be a young adult (Howell 1960; Ovey 1964). The thick parietals appear to have been relatively flat with straight inferior segments and pre-lambdoidal flattening. The maximum breadth is mid-way up the parietals presenting a circular transverse contour in norma occipitalis ("en bombe" form). Biasterionic breadth is quite large. The thick occipital is well rounded with only a slight degree of angulation. There is a moderately developed tripartite transverse occipital torus. The lateral portions are more prominent than the flattened medial segment, above which there is a suprainiac fossa (Hublin 1984; Santa Luca 1978). There is no external occipital protuberance, and the inion is separated from the endinion. It is possible that Swanscombe had prominent occipitomastoid crests (Santa Luca 1978; Stewart 1961b).

Group 3 Predmost 3, Cro-Magnon 1. Group 4 Predmost 4, Mladec 1, Abri Pataud 1.

These results are in general agreement with those of Corruccini (1974a) and Stringer (1974b). Stringer (1978) found Steinheim to be well separated from the Neanderthals, Petralona and Arago 21. Using Mahlanobis' Generalised Distance statistic, Weiner and Campbell ( 1964) found Steinheim to be close to Skhfil and Swanscombe. Hills and Brothwell (1974) found that on an analysis of the frontal squama Steinheim was separated from the Neanderthals, Zuttiyeh, Skhiil 5 and Upper Palaeolithic material, and was closest to anatomically modem hominids. Van Yark (1983, 129

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The date of the Swanscombe deposits is still open to discussion. The hominid remains were recovered from near the base of the Upper Middle Gravels of the 30m terrace of the Thames River (Ovey 1964). The stratigraphical sequence, lithic material, and mammalian fauna from the site have generally been interpreted as indicating a Hoxnian interglacial age equating to either oxygen isotope stage 9 giving an age of around 300,000 years BP or oxygen isotope stage 11 giving an age of around 400,000 years BP {Table 5: 1; Grun et al. 1997; Howell 1960; Oakley 1957; Oakley et al. 1971; Ovey 1964; Stringer 1998; Sutcliffe 1964). However, it has also been suggested that the mammalian remains and molluscan material from the Upper Middle Gravels, and the geomorphological and sediment evidence from the site suggest cooler conditions than those of the Hoxnian interglacial (Bridgland 1980; Bridgland et al 1985; Cook et al. 1982; Kerney 1971; Stringer et al. 1985). This may indicate a late oxygen isotope stage 7 or early stage 6 date, and would provide an age of around 190,000 years BP for the Upper Middle Gravels {Table 5: 1; Griin et al. 1997).

As previously mentioned, there are similarities between the mammalian fauna from Swanscombe and Steinheim (Sutcliffe 1964). Steinheim is usually dated to the Holstein interglacial and given an age of 300-400,000 years BP. Fluorine, uranium and nitrogen analyses indicate that the hominid bones were contemporary with the fauna from the Upper Middle Gravels (Oakley 1980; Oakley and Gardiner 1964: Oakley et al. 1971). Szabo and Collins (1975) attempted to obtained uranium series ages for three samples from Swanscombe, but only one of the samples could be dated. This sample, which came from just below the skull horizon, provided a finite date of 326,000+99,000-54,000 years BP, and ages of greater than 164,000, and 272,000 years BP (Szabo and Collins 1975). Cook et al. (1982:32) feel that problems with the use of this method at the site mean that the results "must be considered a relatively crude estimate of the age of the hominid".

131

Thermoluminescence ages for the site have also been obtained (Bridgland et al. 1985). The Middle Gravel was bracketted between two "apparent" ages of 202,000+/-15,200 years BP, and 228,800+/-23,300 years BP, which place the deposit broadly within the latter half of OIS 7. The dates obtained, however, ranged from approximately 188,000 years BP to 245,000 years BP (Bridgland et al. 1985). Once more a late Middle Pleistocene date for the hominid remains s a reasonable estimate. Swanscombe was only used in the analysis of the occipital squama, where it clustered with La Chaise-Suard 2 and Saccopastore 1 (Fig. 5: 14). There were high loadings on the variables that document the curvature and breadth of the occipital squama (Fig. 5:14). In earlier analyses Swanscombe was a member of the group that included hominids such as Steinheim, Fontechevade 5, Saccopastore 1, Skhiil 5, Jebel Qafzeh 6 and Omo 1 (Habgood 1982, 1984b; Habgood and Walker 1986). Weiner and Campbell (1964) found that according to Penrose's shape distance Swanscombe grouped with La Chapelle 1 and Kabwe 1. When they used Mahlanobis ' Generalised Distance Swanscombe was found to be close to Steinheim , Fontechevade 2, Skhiil 5, Jebel Qafzeh 6, and Tabiin Cl. Stringer (1974b) found that Swanscombe displayed similarities to Vertesszollos 2, Skhiil 5, Fontechevade 2, Omo 1, and early Neanderthals, while on plots of cranial angles and indices he found it to fall closest to Fontechevade 5, Saccopastore 1 and Steinheim, although Kabwe 1, Singa and the Neanderthals were not far removed. Corruccini (1974a) found Swanscombe to be placed in an intermediate position between Homo erectus and Neanderthal calvaria and closest to Steinheim and Kabwe 1. The results are consistent with the total morphological pattern of Swanscombe. Although broader and sagittally longer, Swanscombe is very similar to Steinheim (Breitinger 1964; Howell 1960; Morant 1938; Stringer 1978; Weidenreich 1943b; Wolpoff 1980a, 1980b). There are also similarities in the degree of parietal flattening with Fontechevade 5. Vallois (1949 , 1954) grouped Swanscombe and F ontechevade 5 together in his pre-sapiens group, whereas Weiner and Campbell (1964) did not find a close similarity between the two . Drennan (1956:74) felt that there was "a remarkably close correspondence" in the posterior mid-sagittal and transverse contours of Swanscombe and Kabwe 1. Although they grouped together, Swanscombe has a much rounder occipital contour, less angulation between the occiput and nuchal plane , and a much less flat and horizontally orientated nuchal plane. It also displays simiiarities in the morphology of its occipital torus to Neanderthal crania, in that it has a tripartite torus with a suprainiac fossa (Hublin 1984; Santa Luca 1978). Wolpoff ( 1980b) argues that, although Swanscombe displays less prelambdoidal flattening, it is very similar to Biache. There is also a resemblance to the smaller La Chaise-Suard 2 occipital. Howell (1951) regarded Swanscombe as Neanderthal. Stringer et al. ( 1979) placed their Homo sapiens Grade 1 or 2, along Arago and Montmaurin . Dean et al.

a "progressive" Swanscombe in with Steinheim, (1998) placed

Swanscombe within a pre-Neanderthal evolutionary stage with Bilzingsleben , Vertesszollos , Atapuerca SH site, Steinheim , and Reilingen , which display incipient Neanderthal morphologies In general , Swanscombe can be easily placed within the spectrum of mid- to late Middle Pleistocene European hominids, and in many respects is a pre-Neanderthal.

Atapuerca - Sima de los Huesos The site of Sima de los Huesos [Pit of the bones ](SH) is within the Cueva Mayor cave complex within the Sierra de Atapuerca, northern Spain. (Fig. 5:1). The SH site has yielded almost 3,000 hominid remains belonging to at least 32 individuals, making it one of, if not the largest sample of Middle Pleistocene hominids found (Aguirrie and de Lumley 1977; Arsuaga et al. 1991; Arsuaga et al. 1997). Within the sample there is a prevalence of remains from adolescents and prime-age adults. There are two well-preserved adult crania numbers 5 and 6, an adolescent calvaria - cranium 4, ten well preserved mandibles, a large dental sample, various postcranial elements and numerous isolated cranial, mandibular and postcranial fragments (Aguirre and de Lumley 1977; Arsuaga et al. 1991; Arsuaga et al. 1997). How the hominids accumulated at the site is not clear, with one option being that the bodies were dumped in the chamber as part of mortuary practice, although Gargett ( 1999) argues that the presence of nonhominid remains supports the contention that the location was a "natural trap" and not a ritual burial site. There is a minimum of 13 calvaria within the SH sample (Arsuaga et al. 1997). Whilst the cranial material exhibits a large amount of morphological variability, cranial discrete traits demonstrate less variation, which has been interpreted as indicating that the sample may represent a single biological population (Manzi et al. 2000). The best preserved crania are At Cr-4, At Cr-5 and At Cr- 6. Arsuaga et al. ( 1997) provide a detailed description and comparative study of the cranial material. At Cr-4 (A T-600) is an almost complete adult male calvaria. It has an angular torus and sagittal keeling on frontal and parietals (Arsuaga et al. 1997). At Cr-5 (AT-700) is a complete adult cranium, which can be articulated with mandible AT-888 . It has small calvaria dimensions, but a relatively large projecting facial skeleton . There is sagittal keeling on the anterior frontal squama and on the parietals. At Cr-5 has been identified as either a small male or a female (mandible AT-888 is identified as that of a male). At Cr-6 is a fragmentary cranium of an adolescent male . The crania have relatively thick cranial bones, maximum breadth low down on the vault, parallel vault walls, well-developed double arched supraorbital tori, well-developed mastoid processes and occipital tori with suprainiac fossa., but do not display, lambdoidal flattening and occipital buns (Arsuaga et al. 1997).Cranial capacity varies considerably among these three crania - cranium 4 - l ,390cc, cranium 5 - 1,125cc and cranium 6- l ,220cc (Lorenzo et al. 1998). Whilst the mandibular sample displays a large range of variation, there is a set of features that clearly align with 132

Neanderthal mandibles (Rosas 2001). Mandible At-I, an adult mandible missing most of both ascending rami and all anterior teeth, has a thick and low body that is divergent anteriorly (Aguirrie and de Lumley 1977). The symphyseal region is receding with little development of a mental eminence, the molars are relatively small, especially the 3rd molar, mental foramen are positioned under PM4-Ml and there is a definite retromolar space. Other mandibles have mental foramen under P4 (At-2 & At-301) and Ml (At-300 & At-1775), the most common occurrence and retromolar spaces of varying lengths (At-1 is 9.6mm while At-792 is 18.5mm) (Rosas 2001). These variations may be due to the age of the individuals (a predominance of adolescents) sexual dimorphism or a large of morphological variation. The SH mandibles present a morphological pattern that parallels the Neanderthal pattern (Rosas 2001).

Biache-Saint-Vaast The posterior portion of a hominid cranium, a maxillae fragment containing all six molars, four isolated premolars, an isolated incisor, faunal remains, and lithic material, have been recovered from an open site on a terrace of the Scarpe River, northern France (Fig. 5: 1; Cook et al. 1982; Stringer et al. 1984; Wolpoff 1980b; Vandermeersch 1978). The hominid material is thought to have belonged to an , adolescent female. The remaining vault is small, gracile and quite low, with marked pre-lambdoidal flattening. The rounded occipital protrudes to form a bun, or "chignon". In norma occipita/is the transverse contour is of the "en bombe" form, with the maximum breadth mid-way up the parietals. The occiput is larger than the nuchal plane. The poorlyexpressed occipital torus, which fades laterally, is tripartite, with its most salient portions on either side of a very slight suprainiac fossa (Stringer et al 1984). Wolpoff (1980b:349), however, stated that there is "neither an inferior or a superior sulcus". There is no external occipital protuberance. The occipitomastoid crests are large and project more than the small mastoid processes that angle inwards.

The large dental sample of over 250 permanent and deciduous teeth, representing a minimum of 27 individuals, also demonstrates a large range of variation in dental size (Aguirrie and de Lumley 1977; Bermudez de Castro et al. 2001). The SH sample exhibits a similarity of many dental traits with Neanderthals (Bermudez de Castro et al. 2001)

The rather undiagnostic faunal sample from the site has been used to place the fluviatile deposits, which contained the hominid remains and the lithic material, within an interstadial of the Riss glacial (Cook et al. 1982; Vandermeersch 1978), or at least to the beginning of the last interglacial (Stringer et al. 1984). However, Cook et al. stated that Biache is a prime example of "rissification" in that it has been assigned to this all embracing complex because it does not fit into any better defined stage. (1982:38) Yokoyama ( 1989) reports non-destructive gamma-ray spectrometry ages of 263,000+53,000-37,000 years BP (UTh) and> 175,000 years BP (U-Pa) for Biache. A late Middle Pleistocene age for the Biache hominid is a reasonable estimate until the dating problems are clarified.

Deposits with human remains at the Sima de los Heusos site have normal magnetization that overprints a partially resolved reversed magnetization compatible with one of the reversal events within the Bruhnes Chron (Pares et al. 2000). Yokoyama ( 1989) reports non-destructive gamma-ray spectrometry ages of 320,000+233,000-73,000 years BP (UTh) and > 175,000 years BP (U-Pa) for At-1. electron spin resonance (ESR) and uranium-series analysis on both human and bear remains indicate an age between 200,000 and 300,000 years BP, which is consistent with macro- and microfauna from the deposits. (Falgueres et al. 2001; Pares et al. 2000). On the mandibular non-metric analysis (Fig. 5:16) At-I was placed closest to Arago 2 and Neanderthal mandibles Spy 2, Saint-Cesaire, Guattari-Monte Circeo 2, Shanidar 2 and Tabiin C2.

Biache was not included in any analyses. It does, however, have clear Neanderthal characteristics in its "chignon", prelambdoidal flattening, "en bombe" type transverse contour, tripartite occipital torus, suprainiac fossa, lack of an external occipital protuberance, small mastoid processes and large occipitomastoid crests (Table 5:6; Cook et al. 1982; Hublin 1984; Santa Luca 1978; Smith 1985; Stringer 1985; Stringer et al 1984). There are strong similarities to the larger Swanscombe, and the smaller Steinheim hominids (Stringer et al. 1984; Vandermeersch 1978; Wolpoff 1980a, 1980b), although Biache has more pre-lambdoidal flattening. Biache also displays many similarities with the La Chaise hominid material, especially that from L'Abri Suard (Wolpoff 1980a, 1980b). There are also similarities between Biache and the posterior of La Quina H5. The Biache hominid is generally regarded as a pre-Neanderthal (Ruhlin 1984; Vandermeersch 1978, 1985). Stringer et al. (1979) placed Biache with Krapina and Gibraltar 1 in their Homo sapiens Grade 2 or 3. Dean et al. ( 1998) placed Biache 1 within their Stage 3 early Neanderthal Stage along with material from Ehringsdorf, Krapina, Saccopsatore, La Chaise-Suard and La Chaise Bourgeois-Delaunay.

Arsuaga et al. ( 1997) undertook principal components analysis of both cranial and occipital measurements. On the plot of the first two principal components from the neurocranial analysis, At Cr-4 and 5 were placed between Neanderthal crania and Homo erectus material. At Cr-4 was positioned near the hominids from Petralona and Kabwe, whereas At Cr-5 was positioned between the larger (male) and smaller (female) Neanderthal crania. On the plot of the first two principal components from the occipital analysis, At Cr-4 was placed near the occipital from V ertesszollos, whereas At Cr-5 and At Oc-IV were placed near Steinheim and the smaller Neanderthals Saccopastore 1 and Tabiin 1. The SH hominid sample presents a large range of morphological variation and a mosaic of archaic and Neanderthal features (Aguirrie and de Lumley 1977; Arsuaga et al. 1997; Arsuaga et al. 2001; Lorenzo et al. 1998; Manzi et al. 2000; Rosas 2001 ). The SH hominids have been placed within a pre-Neanderthal evolutionary stage along with Bilzingsleben, V ertesszollos, Swanscombe, Steinheim and Reilingen, which display incipient Neanderthal morphologies (Dean et al. 1998) 133

mention a uranium series age of approximately 245,000 years BP for La Chaise-Suard . A thermoluminescence date of 126,000 +/-15,000 years BP has also been obtained from a burnt stone from La Chaise-Suard. The hominids from La Chaise-Suard need be no earlier than the onset of the last glacial, but it could be earlier.

La Chaise-L'abri Soard L'abri Suard is one of a number of cave sites at La Chaise (see Cook et al. 1982, Debenath 1977, Debenath and Piveteau 1969, Stringer et al. 1984 for the l'abri BourgeoisDelaunay), Charente, central-western France (Oakley et al. 1971). Much of the hominid sample was recovered from deposits between two travertine layers, although the stratigraphical position of the initial collection of hominid remains from the site is unknown . Two of the most complete hominid specimens are La Chaise-Suard 1, a calotte, and La Chaise-Suard 2, an occipital (Cook et al. 1982; Mann and Trinkaus 1973; Piveteau 1957, 1970; Stringer et al. 1984; Wolpoff 1980a, 1980b).

La Chaise-Suard 2 was used in the occipital analyses. It grouped with Swanscombe and Saccopastore 1 (Fig. 5:14) . There were high loadings on the variables for occipital breadth, and occipital curvature (Fig . 5: 14). These results are generally consistent with the morphology of La Chaise-Suard 2, and with earlier metrical analysis of it (Stringer et al. 1984). La Chaise-Suard 2 resembles other late Middle Pleistocene occipitals, especially Biache (W olpoff 1980a, 1980b). It is also pre-empting the later Neanderthals with the configuration of its transverse torus (Table 5:6). In the analyses the closest Neanderthal to it was Saccopastore 1, which is probably also quite close to La Chaise-Suard in date, although Gibraltar 1 (also probably close to La ChaiseSuard chronologically) and La Ferrassie 1 were not markedly removed from La Chaise-Suard 2. La Chaise-Suard 1 displays similarities to Fontechevade 5, Cova Negra, and Neanderthal crania (Mann and Trinkaus 1973; Wolpoff 1980a). The La Chaise-Suard (and also those from Bourgeois-Delaunay) hominid material may be regarded as representing pre-Neanderthals (Cook et al. 1982; Debenath 1977; Hublin 1984; Mann and Trinkaus 1973; Piveteau 1970; Stringer et al. 1984; Vandermeersch 1985). Dean et al. ( 1998) placed the La Chaise-Suard and La Chaise BourgeoisDelaunay remains within their Stage 3 early Neanderthal Stage along with Biache 1 and material from Ehringsdorf , K.rapina and Saccopsatore.

La Chaise-Suard 1 consists of an incomplete parietal and the left half of the frontal squama of a young individual, although Oakley et al. ( 1971) state that the individual was an adult. The metopic suture is preserved . The vault is low and quite thick , while the parietal height is only moderate. The maximum cranial breadth would have been located towards the mid-parietal region . In norma occipitalis the transverse contour would have been circular. The parietal angles are said to resemble Homo erectus more than most other Middle Pleistocen e European Hominids (Wolpoff 1980b). La Chaise-Suard 2 is an almost complete adult occipital. It is relatively small, moderately thick , well rounded in contour, and has a high occipital angle. The highly-curved occiput is larger than the nuchal plane. The transverse torus is not excessively large. It is tripartite in form with the most prominent portions lateral to a very flat medial segment , above which there is a deep suprainiac fossa. The upper margin of the torus is generally not well delineated from the occiput, whereas the lower margin is clearly defined by the excavation of the nuchal plane below it. There is no external occipital protuberance .

Fon tech evade Hominid remains, faunal material and stone artefacts were recovered from a cave near the village of F ontechevade, Charente, central-western France (Fig. 5:2; Boule and Vallois 1957; Day 1977; Henri-Martin 1957; Oakley et al. 1971). The two most important hominid remains from the site are Fontechevade 4 and 5, formerly I and II respectively (Brace 1964; Cook et al. 1982; Henri-Martin 1957; Howell 1957; Stringer et al. 1984; Trinkaus 1973; Vallois 1949, 1954; Vandermeersch et al. 1976).

The dating of the site remains open to some degree of doubt. The sediments , pollen and fauna from La Chaise-Suard are generally assigned to Riss "Ill" , the latter part of oxygen isotope stage 6 (Table 5: 1; Cook et al. 1982; Debenath 1975, 1977; Mann and Trinkaus 1973; Oakley et al. 1971; Stringer et al. 1984; Wolpoff 1980a, 1980b). The lithic material from La Chaise-Suard is classified as Acheulean (Oakley et al. 1971), although there are problems with this attribution (Cook et al. 1982).

Fontechevade 4 is a small frontal fragment consisting of part of the squamous portion and the glabellar region, including the medial section of the left superciliary arch. The fragment is relatively thin and displays little development of the supraorbital region. Vallois (1949 , 1958) felt that the bone thickness, wide interorbital region, and especially the presence of a large frontal sinus indicated that Fontechevade 4 was an adult that lacked well-developed brow-ridges . However , this attribution has been doubted. Trinkaus (1973) has demonstrated that Fontechevade 4 could have been an adolescent Neanderthal aged about 12 years that was still to develop a fully adult brow-ridge. Others have also suggested that the morphology of F ontechevade 4 could be matched by immature Neanderthals (Brace 1964; Howell 1957; Vandermeersch et al. 1976; Wolpoff 1980a). The small size of the fragment , however, precludes any definite conclusions to be made about it.

The lower travertine has produced uranium series dates of 231,000+/-20 ,000 years BP (Cook et al. 1982), and 185,000+/-30,000 years BP (Schwarcz 1980), whereas the upper travertine provided a range of dates from 101,000 to 45,000 years BP (Cook et al. 1982). The upper travertine may correspond with the hominid-bearing bed 11 in La Chaise-Bourgeois -Delaunay which on the basis of pollen counts , was attributed to the last interglacial, but which has provided uranium series ages of 152,000+/-16,000 years BP (Debenath 1977; Cook et al. 1982), and 145,000+/-16,000 years BP (Schwarcz 1980). The early hominid sample may be younger than the base of the upper travertine with a maximum age, based on the uranium series determinations , of 101,000 years BP (Cook et al. 1982). Schwarcz (1982) stated that the hominids from the La Chaise sites range in age from 240,000 to 150,000 years BP. Stringer et al. (1984) 134

Fontechevade 5 is a partial calotte, consisting of the left parietal, the superior section of the right parietal and most of the squamous portion of the frontal. The vault bones are thick, the parietals are flat, biasterionic breadth would have been great, and the junction with the temporal was quite low. In norma verticalis the maximum parietal breadth is posteriorly placed, while in norma occipitalis the maximum vault is located mid-way up the parietals presenting a rounded transverse contour ("en bombe" form). Based on the remaining portion of the frontal squama, Vallo is ( 1949, 1958; Boule and Vallois 1957) concluded that Fontechevade 5 would have had a relatively vertical forehead, little postorbital constriction, and lacked marked brow-ridges. However, this reconstruction has been strongly questioned (Brace 1964; Trinkaus 1973; Vandermeersch et al. 1976; Wolpoff 1980a). Metrical studies have demonstrated that Fontechevade 5 was most similar to Neanderthal or earlier crania (Corruccini 1975b; Sergi 1958; Stringer 1974b, 1978; Weiner and Campbell 1964).

Steinheim. Vallois (1949, 1954) regarded Fontechevade 4 and 5 and Swanscombe as pre-sapiens, and separate from his pre-Neanderthals such as Steinheim, and later Neanderthals. There are similarities between Fontechevade 5 and Swanscombe in the degree of parietal flatness, although Weiner and Campbell (1964) did not find the two to be close. Similarities to Neanderthal crania have already been mentioned (Brace 1964; Drennan 1956; Corruccini 1975b; Howell 1951, 1958; Trinkaus 1973; Stringer 1974b, 1978; Vandermeersch et al. 1976; Wolpoff 1980a). Smith (1982:697) suggests that Fontechevade 5 "in most ways makes a nice fit for the Bilzingsleben occipital", while Wolpoff (1980a) argues there is a similarity in transverse contour with La Chaise-Suard 1. The sagittal curvature of Fontechevade 5 also recalls that of Steinheim. In the past Fontechevade 5 was often seen as displaying a more modem morphology than the Neanderthals, whereas now it is generally seen as presenting a similar or even more archaic morphological pattern.

Generally speaking, the F ontechevade specimens are not complete enough for major conclusions to be made about their morphology, especially in regard to the missing portions.

Howell (1951) thought that Fontechevade 5 may be a "progressive" Neanderthal. Stringer et al. ( 1979) placed Fontechevade in their Homo sapiens Grade 2, along with Saccopastore 1, La Chaise and Ehringsdorf.

The deposits that contained Fontechevade 4 and 5 are generally thought to be either last interglacial or late "Riss" in age (Boule and Vallois 1957; Oakley et al. 1971; Trinkaus 1973; Vandermeersch et al. 1976). The fauna} sample from the site contains both warm and cold elements, and there are similarities to British faunal samples of pre-Eemian and postHoxnian age (Cook et al. 1982). Stringer et al. state ...a consensus seems to be emerging for the attribution of these levels to a "Riss" age. (1984:88) This consensus is based on studies of the microfauna, pollen and Tayacian lithic material (Cook et al. 1982; Stringer et al. 1984).

Ehringsdorf The remains of at least eight hominids, lithic material and a large faunal sample have been recovered from two adjacent travertine quarries (Fischer's and Kampfe's) at Ehringsdorf near Weimar, Germany (Fig. 5:3; Jelinek 1969; Oakley et al. 1971). Ehringsdorf 9 (H), recovered from Fischer's Quarry, is the most complete cranium (Boule and Vallois 1957; Cook et al. 1982; Jelinek 1969; Smith 1984; Stringer et al. 1984; Wolpoff 1980a). This calvaria has been reconstructed on a number of occasions resulting in varying degrees of resemblance to modem or Neanderthal crania (Jelinek 1969). These differences in reconstruction have been especially evident in the slope of the frontal squama, degree of lambdoidal flattening, and the transverse curvature of the vault. These differences have been caused, to a large extent, by the lack of articulation between the vault fragments. Another reconstruction has an angulated occipital squama, a moderately vertical frontal bone, and relatively straight-sided parietals presenting an "en maison" form transverse contour (Stringer et al. 1984).

Fontechevade 4 was not recovered in situ, but was found in a block ofbreccia back in the laboratory (Day 1977; Stringer et al. 1984; Trinkaus 1973). This has led to doubts about its exact stratigraphical placement, and to the suggestion that it may be younger than Fontechevade 5 (Howell 1958; Smith 1982). Fluorine analyses, however, support the attribution of both Fontechevade 4 and 5 to the Tayacian levels (Oakley 1980; Oakley et al. 1971). It should be noted that according to Vallois (1949) the bones had been affected by fire, and this can affect subsequent fluorination of the bone (Oakley 1980).

In general, Ehringsdorf 9 has marked postorbital constriction. The moderately-developed browridges, which are separated from the frontal by an ophryonic groove, do not appear to thin laterally. The supraglabellar segment of the frontal squama is quite steep, and there is a frontal bulge. The highly-flexed occipital squama displays a tripartite occipital torus, with the lateral segments most prominent and with a small suprainiac fossa above the flattened middle portion. The mastoid processes are quite large, although Smith (1984) suggests this is visually accentuated by the occipitomastoid crests being missing.

Fontechevade 4 and 5 were not used in any analyses. In earlier analyses F ontechevade 5 was a member of a group that included Steinheim, Saccopastore 1, Ehringsdorf 9, Swanscombe, Skhul 5, and Omo 1 (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). Stringer (1974b) found Fontechevade 5 not to be significantly different from Neanderthal crania, Saccopastore 1, Petralona, Jebel Irhoud 1, Swanscombe, Kabwe 1 and Omo 1, but to be different to Steinheim, Skhiil 5/9, and especially his Upper Palaeolithic and modern samples. On a plot of parietal indices Stringer (1978) found Fontechevade 5 to be positioned near Saccopastore 1, Kabwe 1 and Singa, and not markedly removed from Steinheim and the Neanderthals. Corruccini ( 1975b) found that Fontechevade 5 had a close affinity to

The parietals recovered from the Ehringsdorf quarries are morphologically quite variable. Parietals 2 (B) and 4 (D) are quite thick and robust, with moderately-developed parietal 135

the upper travertine . Elsewhere Schwarcz ( 1982) suggests that the Ehringsdorf hominids are older than 150,000 years BP, but younger than 250,000 years BP. These dates would place the lower travertine, at least, into the penultimate interglacial and oxygen isotope stage 7, which dates between 190,000 and 245,000 years (Table 5:1; Grun et al. 1997), or within an interstadial of the penultimate ice age. Schwarcz et al. ( 1988) also report electron spin resonance (ESR) age estimates for the upper and lower travertine deposits, which suggest that the upper travertine equates to oxygen isotope stage 5, whereas the lower travertine equates to oxygen isotope stages 6 and 7. Grtin and Stringer ( 1991) supported the uranium-series and ESR age estimates (Grun and Stringer 1991: Fig. 12) and the placement of the site into the penultimate interglacial (oxygen isotope stage 7).

eminences and relatively straight sides, presenting an "en maison" shape in norma occipitalis (Jelinek 1969). Parietal 3 ( C) and those on Ehringsdorf 9 (H), are thinner and more rounded in contour, producing an "en bombe" form in norma occipita/is (Jelinek 1969). Two mandibles have been recovered from Kampfe ' s Quarry (Boule and Vallois 1957; Cook et al. 1982; Jelinek 1969; Oakley et al. 1971; Stringer et al. 1984; Wolpoff 1980a). Ehringsdorf 6 (F) is a relatively complete adult mandible. The corpus is essentially complete , while the left ramus is totally missing and only the anterior portion of the right ramus is preserved. The mandible has a narrow dental arc, although it has been suggested that this may be due to postmortem deformation (Boule and Vallois 1957). There are reduced third molars, retromolar gaps and single mental foramina under the first molars . Pronounced alveolar prognathism and incipient development of the mental eminence are also evident on Ehringsdorf 6 (but see Schwartz and Tattersall 2000) . The mandibular foramen pattern on the right ramus is indistinct (Smith 1978b).

Whilst the revised dates infer an earlier date for the site, the precise date of the Ehringsdorf hominids cannot be confidently ascertained until future research clarifies the dichotomy between the relative and absolute ages for the site. As there are problems in the use of the uranium series method (Cook et al. 1982; Turekian and Bada 1972; Schwarcz 1980), the last interglacial age, based predominantly on the fauna! sample from the site, cannot be completely disregarded . This younger date can also be supported by the lithic material from the lower travertine which is "distinctly Mousterian in nature" (Smith 1984:170; Cook et al. 1982; Coon 1962; Oakley et al. 1971).

The immature mandible, Ehringsdorf 7 (G), consists of the left corpus and ascending rami and the anterior portion of the right corpus. Oakley et al. ( 1971) contend that this mandible is actually two fragments from different mandibles (designated Ehringsdorf 7 and 8) that were incorrectly reconstructed as a single individual. Ehringsdorf 7 (G) has a relatively high, broad and posteriorly angled left ascending ramus, a retromolar gap and incipient development of a chin. Boule and Vallois (1957) and Smith (1984) have stated that the mandible does have an incipient mental eminence. The anterior teeth are large, as is the third molar, and the molars are moderately taurodont. Overall, the Ehringsdorf hominid material displays a great deal of morphological variation .

Measurements were taken on a cast of one of the early reconstructions of Ehringsdorf 9 (Kleinschmidt's reconstruction, cf. Wolpoff 1980a:Fig . 11.5:B). It grouped with the Neanderthal crania Gibraltar 1, La Chapelle 1 and La Ferrassie 1. There were high loadings on the variables that document frontal breadth and curvature of the posterior of the cranium. In earlier analyses Ehringsdorf 9 (W eidenreich's reconstruction , Boule and Vallois 1957:Fig.115) was found to be a member of a group that included Steinheim, Skhul 5, Saccopastore 1, and Gibraltar I (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). As mentioned earlier, due to the lack of articulation of the cranial fragments in all of the reconstructions , their accuracy must remain questionable, as must the results of metrical analyses based on them. Corruccini (1974a) found Ehringsdorf 9 to be most similar to Saccopastore I and then Krapina C.

As with many of the early European hominid sites there is some confusion surrounding the dating of the Ehringsdorf site. The geology, fauna, and flora from the lower travertine at the site, which contained the hominid remains , has generally been taken to indicate a last interglacial age (Boule and Vallois 1957; Oakley et al. 1971). Some questions have been raised about this attribution and especially the fauna! sample, which includes some cold elements. Cook et al. (1982) feel that these variations on a "classic" interglacial assemblage can be incorporated within the complex sequence of the last interglacial as revealed by the oxygen isotope record (Table 5: 1; Grun et al. 1997). They stated The whole sequence could represent the Eemian in the broadest sense, with the lower travertine being the equivalent of substage 5e. (1982:43)

In the analysis of mandibular non -metric traits, Ehringsdorf grouped with the Amud I mandible , and was placed near the Spy 1 and La Quina H5 mandibles (Fig. 5: 16). The intermediate position of Ehringsdorf 6 between Neanderthal and anatomically modem mandibles, was predominantly caused by its incipient development of a mental eminence.

A number of attempts have be made to obtain uranium series dates for the Ehringsdorf travertines (Cook et al. 1982; Schwarcz 1980). The early attempts provided dates of 60,000 to 120,000 years BP, 115,000+/-30,000 years BP, and greater than 220,000 years BP. More recent attempts have provided ages of 225,000+/-26,000 years BP, and greater than 350 ,000 years BP for the lower travertine , and 118,000+/-30,000 years BP (which is regarded as a crude estimate) for the upper travertine (Cook et al. 1982). Schwarcz (1980) provides uranium series dates of 205,000+/-90,000 years BP for the lower travertine, and 146,000+/-30,000 years BP for

The Ehringsdorf hominid material displays morphological affinities to both Neanderthal and earlier hominids . Wolpoff ( 1980a) highlights the archaic features of Ehringsdorf 9 and suggests similarities, especially of the frontal squama, between it and the Steinheim cranium. Coon (1962) suggested similarities between Ehringsdorf 9, Steinheim and Swanscombe. Smith (1984) argues there are marked similarities in features such as the form of the supraorbital region between Ehringsdorf 9 and the Krapina material. Parietals B and D display an "en maison" form with relatively straight sides, while the occipital squama on 136

Ehringsdorf 9 is markedly angulated, which are both archaic features. The more rounded "en bombe" form of the C parietal and those on Ehringsdorf 9 recall Neanderthal parietals, while the occipital bone on Ehringsdorf 9 has a well-developed Neanderthal type transverse torus with a suprainiac fossa.

Arago 47, Swanscombe and Steinheim and features of the occipital that align it with Neanderthals (Dean et al. 1998). On a principal components analysis of five linear measurements, Reilingen was placed closest to Petralona and then European Neanderthals (Swanscombe and Steinheim were not included in the analysis; Dean et al. 1998).

The mandibles, Ehringsdorf 6 and 7 (and 8?), also have the typical Neanderthal traits of the mental foramen positioned under the first molar and retromolar gaps. They do, however, have incipient development of a chin. Smith (1984) argues that the total morphological pattern at Ehringsdorf aligns the hominids with the Neanderthals.

The Reilingen hominid has been equated with both Homo erectus and Neanderthals (see Dean et al. 1998). Dean et al. ( 1998) placed the Reilingen hominid within a preNeanderthal evolutionary stage along with Bilzingsleben, V ertesszollos, Swanscombe, Steinheim and Atapuerca SH, which display incipient Neanderthal morphologies. Such a placement is consistent with the overall morphological pattern of the hominid.

The material does display typical Neanderthal features (Table 5:6), but there are some features, especially evident on the parietals, which sit less easily within the Neanderthal sample. Some of the parietals are clearly different from those of the Neanderthals (Stringer and Trinkaus 1981). Stringer et al. (1984) contend that the parietal morphology may be a wrique derived trait compared with Homo erectus and a shared primitive trait with respect to anatomically modem crania, whereas Smith ( 1984) argues the variation in parietal form at Ehringsdorf can be matched at other Neanderthal sites such as Spy (Stringer and Trinkaus 1981; Thoma 1975). Stringer ( 1978) found that Ehringsdorf 9 has a high frontonasal angle, which separates it from the "classic" Neanderthals.

Upper Pleistocene Hominids This period is taken to commence at around 130,000 years BP, which marks the main transgression of the last interglacial, and to continue until approximately 10,000 years BP when the Holocene period began. In this section, however, only hominids that date in excess of 20,000 to 30,000 years BP will be dealt with in any detail because it is the development of the Neanderthal morphological pattern (Table 5:6) and the subsequent change (transition?) to an anatomically modem morphology that is the focal point of the investigation. Discussions of the hominid remains not described here may be found elsewhere (Boule and Vallois 1957; Brace 1964; Brose and Wolpoff 1971; Coon 1962; Day 1986; Facchini and Giusberti 1992; Green et al. 1981; Howell 1951, 1952, 1957, 1960; Howells 1973a, 1974a, 1974b; Hrdlicka 1927, 1930; Jelinek 1969; Keith 1915, 1925; de Lumley, M-A. 1973, 1975; Mann and Trinkaus 1973; Oakley et al. 1971; Quam et al. 2001; Smith 1982, 1984; Stringer 1984c; Stringer and Currant 1987; Stringer et al. 1984; Wolpoff 1980a).

Howell (1951) and Vallois (1954, 1958; Boule and Vallois 1957) have regarded the Ehringsdorf hominids as "progressive" or pre-Neanderthals". Stringer et al. (1979) placed Ehringsdorf in their Homo sapiens Grade 2, along with Saccopastore 1, Fontechevade, and La Chaise. Dean et al. ( 1998) placed the Ehringsdorf remains within their Stage 3 early Neanderthal Stage along with Biache 1 and material from Krapina, Saccopsatore, La Chaise-Suard and La Chaise Bourgeois-Delaunay.

Reilingen

Alicante

A well-preserved calvaria cons1stmg of two parietals and much of the right temporal and the occipital squama was recovered from a gravel pit near Reilingen, southwestern Germany (Dean et al. 1998).

A stone object resembling a hominid cranium was recovered from among rubble at a commercial excavation site in Alicante, Spain (Fig. 5:1; Cuenca and Walker 1980, 1981). The object is an ectocast of a cranium (lacking a mandible) with four attached cervical vertebral bodies dislocated behind the right mandibular fossa (Cuenca and Walker 1980, 1981). The vault is low and very broad, with a high degree of postorbital constriction. In norma verticalis the maximum breadth is located in a posterior position. The maximum cranial breadth is high up on the parietals. The supraglabellar segment of the well-rounded frontal squama is quite steep. The well-filled out parietals present a smoothly rounded contour in both the sagittal and coronal planes. There is no pre-lambdoidal flattening or occipital protrusion. The occipital squama is well rounded and does not display a transverse torus. The flat, non-prognathic face is quite short, although some of it may have been broken off or eroded away. There is no development of a supraorbital torus, although there are superciliary arches. The glabellar region is not prominent. The interorbital region is quite wide. The high orbits are irregularly shaped, although they do approximate squares. The nasal aperture, which appears to display the

The Reilingen calvaria is relatively low and elongated with maximum breadth at the supramastoid crests and an angled occipital squama presenting an incipient occipital bun. The occipital view of the calvaria presents a pentagonal shape. The parietals display sagittal keeling on the anterior portion, an angular torus and a strongly developed mastoid process. A developed occipital torus and a well-defined suprainiac fossa are present on the occipital. Whilst the date of the hominid cannot be accurately ascertained, it has been related to the oldest fauna from the gravel pit, which ranges from the Holstein interglacial to the Late Wurm glacial (Ziegler and Dean 1998). An amino acid racernization date of 210,000 years BP is reported to have been obtained from a small piece of occipital bone (Dean et al. 1998). The Reilingen calvaria displays features on the parietals that are comparable to archaic Homo sapiens such as Petralona, 137

We have shown the object to be rather unlike Upper or Middle Palaeolithic skulls. (1980:798) Stringer et al. concur with this view and contend that if the Alicante object ...does represent a genuine early Upper Pleistocene fossil, its high rounded cranial shape, without a supraorbital torus, and its anatomically modem facial morphology are quite unprecedented in the European fossil record. (1984:72)

inferior and middle conchae fused to the nasal septum, is imperfectly preserved , but approximates an oval shape . Canine fossae appear to be present. Although the object displays a low cranial vault , the overall morphology is essentially modem. However, there is the problem of the identification of the Alicante object as a fossil ectocast. Cuenca and Walker ( 1980, 1981) discuss the possibility that the object may have been an intentional deception ( a Spanish Piltdown?) or an example of funerary sculpture. They reject these possibilities , but conclude that the identification of the object as a hominid ectocast remains unproven. The object does seem to display specific morphological details such as cervical vertebral bodies , canine fossae, nasal septum and conchae, and external auditory meatus(?), which support the suggestion that it is the ectocast of a hominid . As mentioned before, the object was recovered among rubble and so it was not found in situ . Cuenca and Walker (1980 , 1981) have equated the material forming the object to a geological stratum at the site which they date to the late Middle Pleistocene/early Upper Pleistocene . Although Cuenca and Walker (1980, 1981) suggest that some "human agency" may have initially buried the object, they feel it is unlikely that the object is the result of an Upper Palaeolithic or Epipalaeolithic intrusive burial. However, Stringer et al. stated that .. .it seems much more probable that the specimen was formed during the late Pleistocene or early Holocene ... (1984:72) As the original stratum that supposedly contained the object lies beneath a modem factory, the possibility of the object having been the result of an intrusive burial into late Middle Pleistocene or early Upper Pleistocene deposits cannot be reassessed. There is an absence of cranial sutures and most other anatomical landmarks such as crests or foramina, and the basion is inaccessible due to the presence of the cervical vertebrae (Cuenca and Walker 1980, 1981). This has meant that only a limited number of measurements could be taken, and many of these must be regarded as only reasonable guesses (Cuenca and Walker 1980, 1981). As mentioned above, there is also the possibility that part of the lower face may have been broken off or eroded away which affects some measurements of the splanchnocranium. The Alicante object was not included in any analyses . In earlier analyses it took up one of two positions (Habgood and Walker 1986). When both neurocranial and splanchnocranial measurements were used it clustered with the anatomically modem Upper Palaeolithic crania Cro-Magnon 1 and 3, and Predmost 3 and 4. When only neurocranial data was used the Alicante object clustered with Steinheim, Saccopastore 1, Tabfin 1 and Gibraltar 1. These results suggest that the Alicante object has a modem splanchnocranium (short, flat and non-prognathic face-although some of the lower face may be missing) and a more archaic, though brachycranial, neurocranium (low, broad cranial vault). Cuenca and Walker (1980, 1981) found that on both comparisons of auriculobregmatic height indices and Penrose size and shape distances based on them the Alicante object was situated with pre-Neanderthal material.

The morphological combination displayed by the Alicante object is unusual for a Middle or Upper Pleistocene hominid, although its morphological features can be individually matched on other hominids . This gives some support to the contention that if it is actually a hominid ectocast it may have been an intrusive burial into older layers. However, for the Alicante object to be an Upper or Epi-palaeolithic intrusive burial a 5-6m deep grave would have had to be dug through gravels and solid rock (including a 1.75m thick marine limestone stratum). There is also the problem of explaining how the skull could have turned into a fossil ectocast geologically and in terms of microfossils compatible with the indurated fossil dune stratum. Such a process would only have occurred if the bones were laid in dune sand which, although consolidating, was still loose enough to close around them and fill the nasal and orbital cavities ( Cuenca and Walker 1980; M.J. Walker pers. comm.). At present (and probably for the future) the Alicante object must remain a palaeontological mystery.

Krapina A large collection of hominid remains, fauna! material and Mousterian stone artefacts have been recovered from a rockshelter on the right bank of the Krapinica River near the village of Krapina, northwest of Zagreb, Croatia, (Fig. 5:5; Gorjanovic-Kramberger 1906; Oakley et al. 1971). The hominid remains were recovered from throughout much of the deposit, but the majority came from strata 3 and 4, the "Zona s Homo" or hominid Zone (Gorjanovic-Kramberger 1906; Smith 1976b).

It has

been proposed that the fragmentation and disassociation of the Krapina hominid material may have been caused by cannibalism (Smith 1976b; Ullrich 1978), although this has been contested by Trinkaus (1985a) who contends that the damage pattern can be explained by sediment pressure, postdepositional disturbance, geological processes and excavation. Russell ( 1987a) studied the breakage pattern of the Krapina material and found that it was inconsistent with cannibalism, but supported the contention that the breakage pattern was caused by sedimentary pressure and/or roof falls. Russell ( 1987b) also found that striations on the bones were consistent with postmortem processing of corpses with stone tools. Russell (1987a, 1987b) favours secondary burial of cleaned bones at Krapina and not cannibalism. If correct, Krapina would provide the earliest "solid" evidence for secondary burial (Shipman 1987).

After their morphological comparisons Cuenca and Walker state 138

...the possible transitional status for Krapina A must presently be viewed very cautiously. (1984:152)

The 850 hominid fragments may represent as many as 80-90 individuals (Trinkaus 1978; Wolpoff 1979), although lower estimates have also been suggested (Smith 1976b ). There are many detailed morphological descriptions of the K.rapina hominid material (Brace 1979; Coon 1962; GorjanovicKramberger 1906; Hrdlicka 1930; Jelinek 1969; Kallay 1963, 1970a, 1970b; Molnar and Molnar 1985; Smith 1976b, 1978a, 1982, 1984; Trinkaus 1978; Wolpoff 1978, 1979, 1980a). Only the crania K.rapina A, C, and E, and mandibles Krapina G and J will be discussed in detail here.

Minugh-Purvis (Minugh-Purvis and Radovcic 1991) originally suggested that the morphology of Krapina 1 could be categorised as transitional between Neanderthals and modem Homo sapiens. Subsequently, Minugh-Purvis (1998) did not identify close morphological similarities between K.rapina A and Skhiil 1 and concluded that Krapina A did not provide evidence that modem genes ... had yet arrived on the European continent by 130ka. (1998:350)

Krapina A is the calotte of a juvenile aged approximately 6-8 years old, consisting of the frontal squama, the anterior portions of both parietals and the left temporal (Coon 1962; Gorjanovic-K.ramberger 1906; Hrdlicka 1930; MinughPurvis 1998; Smith 1976b, 1984; Wolpoff 1980a). Coon (1962) suggested the individual was aged 3-5 years, whereas Smith (1976b; Rink et al. 1995) and Minugh-Purvis (1998) propose an age of 6- 8 years. K.rapina A came from stratum 8, one of the higher levels at the site. None of the cranial sutures, including the metopic suture, show evidence of closure, but the bregmatic fontanelle is closed. Smith (1976b) suggested that postmortem deformation resulted in some flattening and broadening of the vault, although this was not mentioned by Minugh-Purvis (1998; Minugh-Purvis and Radovcic 1991). The supraglabellar segment of the frontal squama is quite steep, although it is not as pronounced as on the younger Skhiil I crania (MinughPurvis 1998). The area around the glabella is very slightly developed and there is only a mild indication of a supraorbital torus, although medial and lateral elements are evident. The superior margin of the orbit is rounded as is typical of Neanderthals. K.rapina A displays parietal bossing located superiorly, presenting a rather flattened "box-shape" in posterior profile (Minugh-Purvis 1998:Figure 4). There is also a certain degree of lambdoidal flattening evident on the parietals. The left temporal reveals a moderately projecting mastoid process and well-developed juxta-mastoid eminence.

K.rapina C, which comes from stratum 4, is the most complete and widely referred to of the Krapina remains (Plate 10). The specimen, which is thought to be a young adult female, consists of the upper portions of the splanchnocranium, a large section of the frontal and much of the right side of the vault (Coon 1962; GorjanovicK.ramberger 1906; Hrdlicka 1930; Smith 1976b, 1984; Wolpoff 1980a). The vault was initially reconstructed as being brachycranial (Gorjanovic-K.ramberger 1906), which has been accepted by many authors (Coon 1962; Hrdlicka 1930; Cuenca and Walker 1980). Smith (1976b) has proposed a reconstruction that provides a longer vault measurement for K.rapina C, which makes it mesocranial. The vault displays marked postorbital constriction. On the low frontal there is a moderate frontal bulge and the thick right parietal displays an eminence. Maximum breadth would have been measured relatively high up on the parietals. The moderately long, low and thick right temporal displays many typical Neanderthal features (Table 5:6; Smith 1976b). The mastoid process is small and the occipitomastoid crest is moderately developed. The supraorbital torus is welldeveloped and divided by a supraglabellar depression and there is a very slight depression at glabella. There is little development of an ophryonic groove, and nasion is not depressed. Although there is some medial reduction and flattening, the brow-ridges cannot be clearly separated into a superciliary arch and supraorbital trigone. Smith argues ...there is somewhat more indication of a division in K.rapina C than in other Neanderthals. (1976b:45) The interorbital region is very wide. The frontonasal suture forms an inverted "V", and the intemasal suture angles sharply so that the right nasal bone extends across the top of the left. The frontal sinuses are large and restricted to the supraorbital torus. The large orbits are squarish in shape. The maxillary sinuses are well-developed, and there is no evidence of canine fossae. The relatively broad and gracile face displays mid-facial prognathism.

Coon ( 1962) and Wolpoff ( 1980a, 1992) have stressed modem, possibly transitional, features on K.rapina A. The differentiation of the medial and lateral elements of the supraorbital on K.rapina A does differ from the supraorbital morphology typical of adult and immature Neanderthals. Minugh-Purvis (1998; Minugh 1983), has suggested that brow-ridge development occurred relatively late, ontogenetically, in early Homo sapiens, but earlier in Neanderthals and that K.rapina A was old enough to have a supraorbital form indicative of its adult browridge morphology. This would infer that Krapina A would have had a supraorbital torus with greater differentiation of the medial and lateral elements than any other individual from Krapina. It is worth noting that the mastoid region of K.rapina A displays a moderately projecting mastoid process, a nontypical Neanderthal feature, and a well-developed juxtamastoid eminence, a typical Neanderthal feature.

K.rapina E, which also derives from stratum 4, is the second most complete skull from the site (Gorjanovic-Kramberger 1906; Smith 1976b; Wolpoff 1980a). This calotte, which is thought to be that of a young adult female, consists of much of the frontal, including the supraorbital region and nasal process, the right parietal and a small segment of the left parietal. As with K.rapina C, K.rapina E was also initially reconstructed as brachycranial. Smith (1976b) has again proposed a new reconstruction that makes Krapina E

However, metrical and morphological comparisons by Smith (1976b) did not differentiate K.rapina A from immature Neanderthals. Smith stated In the final analysis, there is no distinct evidence that the A cranium is not Neandertal. (1976b:35) and that 139

mesocranial. The vault is low, small and quite gracile when compared to Krapina C. There is marked postorbital constriction. The supraorbital torus is well-developed and separated from the frontal by a small but distinct ophryonic groove. There is a slight degree of lateral thinning, but there is no mid-toral flattening as on K.rapina C. The frontal sinuses are large and more symmetric than those of Krapina C. They are basically confined to the torus, but do extend into the interorbital region. The form of the frontonasal and intemasal sutures are the same as those of Krapina C. Krapina E has a higher nasal angle than Krapina C.

such as an interglacial (Malez 1970a, 1970b, 1978). Stratigraphical analysis of the fauna indicates two warm periods and two cold periods (Malez 1970b). Malez (1970a, 1970b, 1978) proposes that Krapina was inhabited from towards the end of the last interglacial to the middle of the "Wurm II" stadia! (following Valoch 1968). Malez (1970a) places strata 1-4 within the late "Riss-Wurm" interglacial, strata 5-7 in "Wurm I", strata -8 in "Wurm 1/11'',and strata 9 in "Wiirm II'' (but see Rink et al. 1995). Smith ( 1976b) contends that Krapina was no longer inhabited by 30,000 years BP. Smith bases this, to a large extent, on the nearby sites of V elika Pecina at which an Aurignacian level has provided a date of 33,850+/-520 years BP (GrN-4979), and Veternica where a date of greater than 43,200 years BP (GrN-4984) has been obtained for a Mousterian level (Smith 1976a, 1982, 1984; Vogel and Waterbolk 1972, but see Smith et al. 1999). A radiocarbon date of 30,700+/-750 years BP (GrN-4299) has been reported for Krapina (Vogel and Waterbolk 1972), but the exact level from which it was obtained is not known (Smith 1976b). Wolpoff (1980a) suggests that Krapina may have been recovered from at, or just below, the level in which this date was obtained .

Mandible G consists of the right corpus and symphyseal region (Gorjanovic-Kramberger 1906; Hrdlicka 1930; Jelinek 1969; Smith 1976b). This adult mandible was recovered from stratum 6. Only a small section of the anterior base of the right ascending ramus is preserved. The three right molars are also present. The inferior and superior edges of the corpus are relatively parallel. There is a slight retromolar gap and a very retreating symphyseal region. The multiple mental foramina are positioned under the right first molar . The gonial angle appears to have been rounded. There is no hypoplasia on the molars (Molnar and Molnar 1985). A well-marked gradient of supraradicular taurodontism from the first to third right malars is evident (Kallay 1963).

Smith (1976b, 1984) cautions that it is possible that the early warm period identified at Krapina and related to the last interglacial may represent an interstadial within the early Wurm (Butzer 1971; Zeuner 1959).

Mandible J (I) is an almost complete (GorjanovicKramberger 1906; Hrdlicka 1930; Jelinek 1969; Smith 1976b). This adult mandible, which was found in stratum 4, is the largest and most well known of the Krapina mandibles. The dental arcade is broad and parabolic in shape. The corpus is long, high and thick. The symphyseal region does not display a well-developed mental eminence. The relatively broad ascending rami are robust and posteriorly angled. The coronoid processes are large and project above the condyles. The sigmoid notches are broad and quite deep. The large, broad and flat condyles have been affected by osteoarthritis. The horizontal-oval (H-O) mandibular foramen pattern (Kallay 1970a; Smith 1976b, 1978b) is evident on the ascending rami. There is a definite retromolar space, and the mental foramina (2 on the left side, 1 on the right) are positioned under the first molars. Linear hypoplasia is evident on both canines (Molnar and Molnar 1985). The left second molar and the right third molar display radicular taurodontism (Kallay 1963).

Electron spin resonance (ESR) and uranium-series dates have more recently been obtained on tooth enamel from the site (Rink et al. 1995). The U-series and early uptake ESR dates were more comparable. A mean age or 130,000 years BP is attributed to layers 1-8, which would place the deposits within the last interglacial (Rink et al. 1995). This mean age for most of the deposit is not consistent with Malez stratigraphical reanalysis of the fauna detailed above. Only skulls C and E and mandibles G and J were used in analyses. Krapina C was found to cluster with the less "classic" Neanderthals Saccopastore 1 and Gibraltar 1, and then with the immature Neanderthal crania La Quina H18 and Teshik-Tash (Figs. 5:12, 5:13, Table 5:5). There were high loadings on the variables that document cranial height and facial size (Fig. 5: 12, Table A: 12). Krapina E joined the Krapina C, Saccopastore 1 and Gibraltar 1 group. In earlier

Within the Krapina skeletal sample there is a great deal of morphological variation, which is contributed to by the inclusion of many subadult individuals, chronological separation, and a high degree of sexual dimorphism. In general, the cranial and postcranial material displays Neanderthal morphological features (Coon 1962; Gorjanovic-Kramberger 1906; Hrdlicka 1930; Jelinek 1969; Kallay 1970a; Smith 1976b, 1978a, 1982, 1984; Trinkaus 1978c; Wolpoff 1980a). The dental remains fit within the Neanderthal sample in size and frequency of taurodontism of the molars (Kallay 1963, 1970b; Smith 1976b; Wolpoff 1978, 1979, 1980a). It is worth noting that no distinct evolutionary trends could be identified within the Krapina hominid sample which may span 50,000 years (Smith 1976b).

Table 5:5. K-means cluster analysis with Guattari-Monte Circeo 1, Saccopastore 1, Krapina C, La Quina Hl8 and Teshik-Tash. Analysis includes 75.1% of the total within group variance. Group 1 Steinheim , Kabwe 1, Gibraltar 1, La Chapelle 1, La Ferrassie 1, Saccopastore 1, Monte Circeo 1, Krapina C, La Quina H18, Teshik-Tash. Group 2 Skhul 5, Combe Capelle, Mladec 1, Chancelade , Abri Pataud 1, Dolni Vestonice 3, Oberkassel 1, Oberkassel 2.

Krapina has usually been dated to the last interglacial (Coon 1962; Oakley et al. 1971) . Re-interpretations of the site have demonstrated that it does not represent a single time period

Group 3 Predmost 3, Predmost 4, Cro-Magnon 1.

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analyses Krapina C was in the same group as Saccopastore 1 and Gibraltar 1, Steinheim, Skhiil 5 and Tabiin CI (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). Measurements of the initial reconstruction of Krapina D (Coon 1962; Gorjanovic-Kramberger 1906) were also used in these earlier analyses (Habgood 1982, 1984a, 1984b; Habgood and Walker 1986). It was found to group with K.rapina C. However, it has since been demonstrated that this reconstruction was incorrect in that fragments from two different individuals had been incorporated into it (Radovcic and Caspari 1987).

The face of the Krapina Neandertals as expressed by the morphology of Krapina C is typically Neandertal in most respects. (1976b:51) and that ...the female Krapina C cranium compares almost exactly to the small, probably female cranium from Gibraltar. This similarity is striking both morphologically and metrically. ( 1978a:108) However , the K.rapina cranial sample does not document the degree of occipital bunning or mid-facial prognathism found on western European Neanderthal crania, and their mid-facial regions are not as robust (Smith 1976b, 1978a, 1982, 1984). K.rapina B, a juvenile from stratum 4, does display some lamdoidal flattening, but fragmentary occipital material and Krapina D do not display flattening near lambda (Smith 1976b). Even though there are these differences between the K.rapina sample and western European Neanderthals Smith argued that the Krapina sample ...do not, as a group, exhibit any feature or complex of features that justifies their being considered "progressive" in comparison with W estem European specimens. (1982:673)

Stringer (1974b) found Krapina C to be nearer his Upper Palaeolithic sample than to the "classic" Neanderthals. Corruccini (1974a) found Krapina C to be similar to Ehringsdorf and Saccopastore 1, and to be the closest European cranium to the western Asian Neanderthals and the hominids from Skhiil. Krapina C is morphologically and metrically very similar to Saccopastore 1 and Gibraltar 1 (Plates IO and 11; Corruccini 1974a; Habgood 1982, 1984a, 1984b; Habgood and Walker 1986; Smith 1976b, 1978a, 1982, 1984; Stringer 1974b; Stringer et al. 1984). Krapina G and J were used in the analysis of mandibular non-metric traits where they were found not to group together (Fig. 5: 11). Krapina J clustered with the "classic" Neanderthal mandibles La Ferrassie 1, Regourdou, Tabiin Cl and Shanidar 1. Other "classic" Neanderthal mandibles (Shanidar 2 and Spy 2) were located nearby. Krapina G was separated from most of the other Neanderthal mandibles. It was part of a group of mandibles positioned between the "classic" Neanderthal cluster and a cluster of mandibles that pre- and postdate most of the "classic" Neanderthals and which are generally thought to display primitive and/or transitional features. This group was made up of Mauer , Malamaud 1, Hortus 4 and La Naulette 1. Near K.rapina G were mandibles that are generally referred to as Neanderthallike in that they display variations on the typical Neanderthal pattern (Table 5:6; Stringer et al. 1984). These mandibles were Arago 3, Bafiolas and Guattari-Monte Circeo 3.

Stringer et al. (1979) placed K.rapina in their Homo sapiens Grade or 3, along with Gibraltar 1 and Biache. Dean et al. ( 1998) placed the Krapina remains within their Stage 3 early Neanderthal Stage along with Biache 1 and material from Ehringsdorf , Saccopsatore, La Chaise-Suard and La Chaise Bourgeois-Delauna y.

Saccopastore Two hominid crania were recovered from a gravel pit on the lower terrace of the Aniene tributary of the Tiber River near Rome, Italy (Fig. 5: 1; Boule and Vallois 1957; Oakley et al. 1971; Sergi 1929, 1931, 1948a). Saccopastore 1 is the almost complete cranium of what is thought to be an adult female (Sergi 1948a, 1948b, 1958; Stringer et al. 1984). The supraorbital region was apparently broken off and lost during excavation. The vault is quite small and low with marked postorbital constriction. The supraglabellar segment of the frontal squama is quite steep. The parietals are quite long and flat with some prelambdoidal flattening. Maximum cranial breadth is mid-way up the parietals , which forms an "en bombe" type transverse contour in norma occipitalis. The occipital is evenly rounded and does not form a Neanderthal "chignon". There is a tripartite transverse occipital torus with a suprainiac fossa above the flattened medial segment (Santa Luca 1978). There are eleven Wormian bones in the lambdoidal region. The mastoid processes are small and there are occipitornastoid crests, but anterior mastoid tubercles appear to be absent (Santa Luca 1978; Stringer et al. 1984). Basicranial flexion is said to be greater than is typical of "classic" Neanderthals (Sergi 1948b). There is both total and mid-facial prognathism . The latter appears to be somewhat less than is common for "classic" Neanderthal crania, while the former is greater (Stringer 1978; Stringer and Trinkaus 1981). The complex frontal sinuses are large and laterally extensive. The

Analyses of non-metric dental traits have found that the Krapina dental sample is different to that of modem human groups (Stringer et al. 1997; Tyrrell and Chamberlain 1998). The results from the analyses are in agreement with the overall morphology pattern of the Krapina hominids in that they do display typical Neanderthal features (Table 5:6; Smith 1976b, 1978a, 1982, 1984). Smith states The total morphological pattern of the Krapina hominids clearly aligns them with Neandertals, and although there is considerable variation, no feature excludes any Krapina specimen from this hominid group ... (1984:145) and that ...the K.rapina Neandertals are not qualitatively separable from Western European Wiirm Neandertals on the basis of any morphological feature. (1982:673) In relation to Krapina C Smith suggests 142

relatively large orbits are circular in shape. The nasal bones are quite prominent, and the large nasal aperture is wide and low. The malars have a high degree of pneumatisation, little indication of canine fossae and display more angulation when they join the maxillae than is common for "classic" Neanderthals . Overall, Saccopastore 1 displays many typical Neanderthal features (Table 5:6; Boule and Vallois 1957; Sergi 1948a, 1948b; Stringer et al. 1984; Santa Luca 1978).

process (Stringer et al. 1984). As with Saccopastore 1, Saccopastore 2 displays numerous typical Neanderthal features (Table 5:6; Boule and Vallois 1957; Sergi 1948a, 1948b Stringer et al. 1984). An age of approximately 60,000 years BP has been quoted for the Saccopastore site ( Oakley et al. 1971; Wolpoff 1980a) . The date of 58,000+/-500 years BP (GrN-2572) was obtained at another site and correlated with the deposits at Saccopastore (Oakley et al. 1971; Vogel and Zagwijn 1967). However, the faunal, floral and molluscan remains from the site suggest a last interglacial age for the Saccopastore hominids (Oakley et al. 1971; Sergi 1948a; Stringer 1982; Stringer et al. 1984). Fluorine and nitrogen analyses demonstrate that Saccopastore 1 and 2 are contemporary with hippopotamus dentine from the same deposit (Oakley 1980; Oakley et al. 1971).

Saccopastore 2 is less complete, larger, and more robust than Saccopastore 1 (Sergi 1948a, 1948b, 1958; Stringer et al. 1984). This adult male skull consists of most of the face, except the left orbital region, and much of the inferior portion of the right side of the vault and cranial base . The large face displays total and mid-facial prognathism, with the former greater and the latter less than is usual for "classic" Neanderthal crania (Stringer and Trinkaus 1981). The right brow-ridge is well -developed . It follows the curve of the large, round, right orbit, and thins laterally . The interorbital region is broad. The ovoid nasal aperture is large. The malar region is similar to that on Saccopastore 1 in that it has expanded maxillary sinuses, little indication of canine fossae and some angulation where the malar and maxillae converge. The cranial base is said to be more flexed than most "classic" Neanderthals (Sergi 1948b), although Laitman et al. (1979) found it to be similar to La Ferrassie 1 and Guattari-Monte Circeo 1. On the plot of a discriminant analysis of the basicranial line, Laitman et al. ( 1979) did not find the Neanderthal crania to form a group, but instead they displayed a high degree of variation. The occipitomastoid crest is only moderately developed, and there does not appear to be an anterior tubercle on the short broad right mastoid

Saccopastore 1 is one of the volatile individuals that moves between groups. It generally clustered with the less "classic" Neanderthal crania Gibraltar 1 and Krapina C (Figs. 5:7, 5:8, 5:12, 5:13). It also grouped with the immature Neanderthal crania La Quina Hl8 and Teshik-Tash (Figs. 5:12, 5:13). On other occasions Saccopastore 1 clustered with Steinheim , Kabwe 1, Petralona and Skhiil 5 (Figs. 2:5, 2:6). When the largest number of variables were used there were high loadings on the variables which describe the curvature of the posterior vault and the size of the face (Fig. 5:8, Table A: 10). In the occipital analyses Saccopastore 1 grouped with Swanscombe and La Chaise-Suard 2 (Fig. 5:14). Saccopastore 2 clustered with Saccopastore 1 and Gibraltar 1.

A2

2

K1.

sw• IOC

Sa1

· ·ov3

.ASB

LF1. •G1

.SL

• •LaC2

0

•LIA -1

.Lie

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'---------------,,-------.-2

- 1

0

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Figure 5:14. Correspondence analysis of the occipital squama. 143

Al

Stringer (1974b) found Saccopastore 1 to display similarities to Skhul 5, Jebel Irhoud 1, Fontechevade 2, Vertesszollos 2, and Omo 1 and 2. In an analysis of the vault using Penrose distances Stringer (1978) found Saccopastore 1 to be closer to the Jebel Qafzeh reference group than other Neanderthals. On an analysis of the face, Saccopastore 1 was closer to Arago 21, the reference hominid, than other Neanderthals (Stringer 1978). When the Neanderthals were used as the reference group for the Penrose distance analyses, Saccopastore 1 was quite close to them. When Stringer ( 1978) plotted cranial angles and indices, Saccopastore 1 was generally closer to the earlier hominids than were other Neanderthals. Stringer (1978) found that the zygomaxillare angle of Saccopastore 2 would have placed it in an intermediate position between the "classic" Neanderthals and earlier hominids such as Petralona and Steinheim. Corruccini (1974a) found Saccopastore 1 to display similarities to Ehringsdorf, and to a lesser extent Krapina C and Steinheim.

1962; Howell 1951, 1952, 1957; Jelinek 1969; Morant 1927; Smith 1982, 1984; Stringer et al. 1984; Wolpoff 1980a). The German site of Salzgitter-Lebenstedt has produced the most northerly Neanderthal so far recovered (Butzer 1971; Hublin 1984; Stringer et al. 1984, but see Pavlov et al. 2001). The Neanderthal sample spans the period from the beginning of the last glacial-oxygen isotope stage 4, which dates to approximately 75,000 years BP, down to at least the end of oxygen isotope stage 3 during "Wurm III", which dates to approximately 30,000 years BP (Table 5:1; Butzer 1981; Gamble 1986a; Griin and Stringer 1991; Laville et al. 1980). The dates towards the end of this range have been recorded for the Neanderthal remains at the sites of Zafarraya and Sima de las Palomas (M.J. Walker pers. comm.), Spain, Saint-Cesaire, France and Vindija , Croatia and even later dates for Mousterian and Chatelperronian occupation layers at sites, especially in Spain and Portugal (Hublin et al. 1995; Mellars 1998a; Mercier et al. 1991; Smith et al. 1999).

The results are in agreement with the morphology of the Saccopastore hominids, for they demonstrate that the two crania have a predominance of Neanderthal features, but in some features they deviate rather from the characteristic "classic" Neanderthal pattern. They have more total and less mid-facial prognathism, less inflated malars, more angulation between the malars and maxillae and possibly more basicranial flexion than is typical of "classic" Neanderthals. This deviation has been highlighted in previous studies of the Saccopastore hominids.

Western Europe has provided a large sample of quite complete Neanderthal cranial and postcranial remains, possible due to the practice of inhumation (Stringer et al. 1984). In eastern Europe burial of the dead does not appear to have been commonly practiced during the time of the Neanderthals (see discussion of the earlier Krapina hominid remains) which has resulted in a very fragmentary Neanderthal sample (Jelinek 1969; Smith 1982, 1984). Some of the fragmentation at eastern European sites may be the result of post-mortem processing of corpses, although evidence for such activity has been suggested at western European sites as well (Defleur et al. 1999). Gargett (1989, 1999) has argued on geomorphological and sedimentological grounds that there is no conclusive evidence for Middle Palaeolithic burials in Europe (but see the comments to Gargett 1989).

Sergi (1948a, 1948b, 1958) felt there was a clear difference between the last interglacial (Saccopastore) and last glacial (Guattari-Monte Circeo) inhabitants of Italy. He stated that Saccopastore I resembles Gibraltar, but differs from the Neanderthalians, including Circeo ... (l 948a:63) Sergi (1948b, 1958) also identified similarities between the Saccopastore crania and the hominids from Krapina and Ehringsdorf, which also probably date to the last interglacial or early last glacial. Stringer ( 1978) felt that this material links the later "classic" Neanderthals with the earlier European hominids. Saccopastore 1 does display morphological and metrical similarities to Krapina C and Gibraltar 1 (Plates 10 and 11; Corruccini 1974a; Habgood 1982, 1984a, 1984b; Habgood and Walker 1986; Sergi 1948b, 1958). Howell (1951) and Vallois (1954, 1958; Boule and Vallois 1957) regarded the Saccopastore hominids as "progressive" or "pre-Neanderthals". Stringer et al. (1984) make more of the similarities that Saccopastore 1 and 2 have to the "classic" Neanderthals and suggest that the differences that are present are primitive retentions. Stringer et al. ( 1979) placed Saccopastore 1 with La Chaise, Ehringsdorf and Fontechevade in their Homo sapiens Grade 2. Dean et al. (1998) placed the Saccopsatore hominids within their Stage 3 early Neanderthal Stage along with Biache 1 and material from Ehringsdorf, Krapina, La ChaiseSuard and La Chaise Bourgeois-Delaunay.

"Classic" Neanderthals Hominids that may be referred to as "classic" Neanderthals have been recovered from throughout much of Europe (Figs. 5:1, 5:2, 5:3, 5:4, 5:5, 5:6; Boule and Vallois 1957; Coon

Most of the relatively complete Neanderthal skeletal material, which is generally associated with industries belonging to either the F errassie or Quina Mousterian groups (Oakley et al. 1971; Vandermeersch 1965), appears to be from the "Wurm II" stadia! and dates around 50,000 to 60,000 years BP. However, Mellars (1986, 1989a, 1989b), based on his contention (Mellars 1969) that there is a sequential development of the Mousterian variants identified by Bordes ( 1968) and thermoluminescence dating of Le Moustier cave (Valladas et al. 1986), gives the La Ferrassie hominids a much earlier date, but this suggestion needs to be further substantiated (Ashton and Cook 1986; Stringer et al. 1984; see also Griin and Stringer 1991 ). There is a suite of features that typify the Neanderthals and generally distinguish them from contemporary hominids from other geographical regions (Table 5:6; cf. Boule 1913; Heim 1974; 1976; Hublin 1984; Trinkaus 1983a; Santa Luca 1978; Stringer 1985; Stringer et al. 1984; Smith 1976b; Wolpoff 1980a). Most of these features are not unique, derived traits for the Neanderthals, in that they can be found on other crania. For example, Kabwe 1 displays mid-facial prognathism (Stringer 1978), the occipital of Eliye Springs 11693 has a suprainiac fossa (Brauer and Leakey 1986a, 1986b), and north African Mesolithic crania have welldeveloped occipital buns (Anderson 1968; Greene and Armelagos 1972; Vallois 1952). However , the combination 144

Table 5:6. Some characteristic features of the cranium and mandible evident on the Neanderthals (cf. Heim I 974, 1976; Hubl in I 984, 1998; Hub Jin et al. 1996; Rak 1998; Smith 1976b, 1978b, 1983; Smith and Ranyard 1980; Stringer 1985, 1998; Stringer et al. 1984; Thoma 1965; Trinkaus 1983a; Wolpoff 1980a; Vandenneersch 1985) . The features listed below are characteristic of the Neanderthals as a whole, but are not necessarily found on every individual. NEUROCRANIUM Large cranial capacity. Long, low and relatively thick vault with marked postorbital constriction . A receding frontal squama . Long and flat parietals . Lambdoidal flattening . Maximum cranial breadth positioned in the mid-parietal region. An occipital bun or chignon with a highly curved lambda-inion segment and a relatively horizontal nuchal plane. Long and low temporal squamae . In norma occipitalis "en bombe" transverse contour. Tripartite occipital tori that are virtually horizontal and of uniform thickness . Distinct suprainiac fossa positioned above the flattened medial segment of the occipital torus. Non-projecting mastoid processes. Large occipitomastoid crests that project below the mastoid processes. Digastric fossae that are closed anteriorly. Large mastoid crests that occupy a high position and end in anterior mastoid tubercles . Supramastoid sulci that narrow anteriorly. The anteroinferior-posterosuperior orientated (elliptical shape) external auditory meatus is positioned superiorly and is in line with the long axis of the zygomatic arch. Shape of the semi-circular canals of the inner ear (Bony labyrinth morphology). The zygomatic root is posterior to the second molar . Horizontally oriented tympanic plate with heavy anterior and posterior portions. Wide and shallow glenoid fossae. A relatively large sphenoidal angle and flattened cranial base. SPLANCHNOCRANJUM Double arched supraorbital torus with the superciliary arch fused with the supraorbital trigone and separated from the frontal by an ophryonic groove and supraglabellar depression. Prominent lateral orbital pillars . Extensive pneumatisation of the supraorbital torus with the frontal sinus extending laterally but not superiorly . Marked mid-facial prognathism. Large upper face relative to facial breadth . Large round orbits . Wide interorbital region. Anteriorly placed and voluminous nasal aperture with projecting nasal bones and a lowered sloping nasal floor. Large infraorbital foramen . Shelving of the malars into the maxillae with little demarcation between the two bones . Retreating zygomatic profile . Extensive pneumatisation of the maxillary sinuses. No canine fossae . Low subspinale angle. Low nasiofrontal angle . Broad anterior palate with bicanine expansion. Dentition positioned anteriorly . Large anterior teeth often with significant attrition and microwear . Incisors with markedly-developed marginal ridges (shovel-shaped) and basal tubercles . MANDIBLE Dentition anteriorly positioned . The dental arcade is anteriorly flattened with bicanine expansion . Large anterior teeth. Taurodontism of the molars. Moderately broad and posteriorly angled ascending rami. Large flat condyles . Rounded gonial angle. Horizontal-oval pattern of mandibular foramen common . Sloping symphyseal region lacking a well-developed mental eminence . Retromolar gap . Multiple mental foramina . Mental foramina positioned under the first molar . Shallow mandibular notch with deepest point near condyle. Coronoid process larger than the condylar process.

of the features is only found on Neanderthal crania and mandibles (Table 5:6). Most of the Neanderthal remains have been described and discussed in detail elsewhere (Boule and Vallois 1957; Heim 1974, 1981-82a, 1981-82b, 1982a, 1982b, 1983; Howell 1951, 1952, 1957; Jelinek

1969; Martin 1923; Morant 1927; Stringer et al. 1984; Smith 1982; 1984; Trinkaus and Howells 1979; and references therein), and so this will not be done again here. Only two sites (Vindija and Saint-Cesaire) that are essential to an understanding of the change (transition or 145

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Figure 5:15. Correspondence analysis of cranial non-metric traits .

• LOS

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BC2 Ch 02 CMl • Pal C M3 DV3

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Figure 5:16. Correspondence analysis of mandibular non-metric traits.

146

replacement) from Neanderthals to anatomically modem hominids will be dealt with in any detail. Other Neanderthal remains will be mentioned when they are relevant to the discussion.

Stringer and Trinkaus 1981; Turb6n et al. 1997) also found on multivariate analyses a clear distinction between the Neanderthals and anatomically modem humans. The Neanderthals, therefore, form a consistent group that is easily distinguished from anatomically modem material. The Neanderthals are shown to have a closer relationship with the earlier hominids, especially Steinheim (Fig. 4:6), than with the contemporary Skhiil (except Skhiil 4) and Jebel Qafzeh hominids or later anatomically modem material (for example Figs. 4:8, 5:8, 5: 14, Table 5:5).

The European Wtirm Neanderthal crania (Plates 10, 11, 12 and 13; Gibraltar 1, La Chapelle 1, La Ferrassie 1, La Quina 5 and HIS, Guattari-Monte Circeo 1, Neanderthal-Feldhofer 1, Spy 1 and 2, Engis 2 and Saint-Cesaire) were found to form a consistent group (Figs. 5:8, 5:9, 5:10, 5:11, 5:12, 5:13, Tables 5:3, 5:4, 5:5). There were generally high loadings on the variables that describe the sagittal and coronal contours of the vault, temporal shape, and facial size (Figs. 5:8, 5:11, 5:13, Tables 5:4, 5:4, 5:5). The European Neanderthal crania also grouped with the western Asian Neanderthal crania (Figs. 4:5, 4:6, 4:7, 4:8, Tables A:6, A:7). In the analyses of the occipital squama the European "Wurm" Neanderthals (Gibraltar 1, La Chapelle, La Ferrassie 1 and Salzgitter-Lebenstedt) were separated from the other material and formed a broad group (Fig. 5:14). In the analysis of non-metric cranial traits there was again a European "Wurm" Neanderthal (Gibraltar 1, La Chapelle 1, La Ferrassie 1, La Quina H5, and Spy 1 and 2) group (Fig. 5:15). These crania were grouped together because they all have occipitomastoid crests, tripartite occipital tori and suprainiac fossa (Fig. 5:15; Santa Luca 1978).

Stringer et al. ( 1979) placed the European "Wurm" Neanderthals, along with the western Asian Neanderthals, into their Homo sapiens Grade 3a. Gibraltar 1 is placed in the Homo sapiens Grade 2 or 3 with Biache and Krapina. Dean et al. (1998) placed Feldhofer, Spy, Monte Circeo, Gibraltar 1, La Quina, La Chapelle-aux-saints, La Ferrassie, Le Moustier (and material from Shanidar and Amud) into their "classic" Neanderthal Stage 4. Coon suggested that the Neanderthals ...are in fact so homogeneous that a strong selective agency must have been pruning off deviant individuals. (1962:529) Although the Neanderthals display a particular combination of characteristic features, they do not display, as Coon suggests, a static morphological pattern. There is quite a lot of morphological variation ( on a particular Neanderthal theme) within the Neanderthal sample (Wolpoff 1980a).

In the analyses of mandibular non-metric traits there is some variation evident with the sample of Neanderthal mandibles (Fig 5:16). There still is a major European and western Asian Neanderthal group in which all the mandibles have multiple mental foramina positioned under the first molars, large retromolar gaps, and lack well-developed mental eminences (Fig. 5:16). Other Neanderthal mandibles, although separated from it, are positioned near the major Neanderthal group, and so combine to form a larger Neanderthal cluster (Fig. 5:16). Positioned away from this cluster was another set of Neanderthal mandibles (La Quina H5, Spy 1, Ehringsdorf 6 and Amud 1). These four mandibles generally have posteriorly positioned, single mental foramina, retromolar gaps and lack well-developed mental eminences (Fig. 5:16). Guattari-Monte Circeo 2 and 3 are well separated from the other "Wurm" Neanderthal mandibles in this analysis (Fig. 5:11; discussed below).

The Gibraltar 1 (Forbes' Quarry) cranium, although having its closest affinities with the "classic" Neanderthals (Trinkaus 1984b; Santa Luca 1978; Stringer et al. 1984), displays a number of differences (Corruccini 1974a; Keith 1925; Habgood 1982, 1984a, 1984b, 1985a; Stringer 1978). In one analysis Gibraltar 1 clustered with Steinheim, Jebel Qafzeh 6 and Skhiil 9 and then Kabwe 1 and Jebel Irhoud 1, before it joined with the "classic" Neanderthals (Fig. 4:9). Gibraltar 1 has a steeper supraglabellar segment of the frontal squama, reduced brow-ridge development, more basicranial flexion (Laitman et al. [ 1979] suggest that the reconstruction may be incorrect), and less occipital bunning than the "classic" Neanderthals (Plate 11). The occipital torus lacks the tripartite configuration of the "classic" Neanderthals, although there is a suprainiac fossa. Weidenreich (1943b) and Condemi (1983) also noted differences in the sphenoid bone between Gibraltar 1 and "classic" Neanderthals.

In all of the analyses the European "Wurm" Neanderthal material was well separated from the anatomically modem Upper Palaeolithic material, Skhiil 5 and the Jebel Qafzeh hominids (Plates 4, 5, 6 and 7). There are no sub-Saharan African, east Asian or Australasian Neanderthals. Although Jebel Irhoud 1 (Plate 3) grouped with the Neanderthals, due to its archaic vault shape, it clearly does not display the characteristic Neanderthal features of the face or occipital (Chapter 3; Santa Luca 1978). El-'Aliya 1 also recalls the Neanderthals in lacking a canine fossa, but there are problems with its dating and the lack of a canine fossa may be a primitive retention in north Africa in that the Rabat maxilla also lacks one (Chapter 3).

The differences between Gibraltar 1 and the other Neanderthal crania may be attributed to sexual dimorphism, with the former being a female, and La Chapelle 1, La Ferrassie 1 and Guattari-Monte Circeo males (Plate 11). The differences may also be caused by temporal separation, with Gibraltar 1 being earlier than the others. Gibraltar 1 was discovered in 1848 by labourers at a limestone quarry, and so it has no stratigraphical, faunal or archaeological contexts (Hrdlicka 1930; Keith 1925; Stringer et al. 1984). Cuenca and Walker ( 1980, 1981) suggested that the brecciated talus that probably contained the cranium (Duckworth 1911; Keith 1925) may be of last interglacial age. This date is consistent with the morphological similarities between Gibraltar I and Krapina C and Saccopastore 1 (Fig. 5: 12, Plates 10 and 11;

Howells (1970, 1989, 1995), Bilsborough (1972, 1973), Andrews and Williams (1973), Corruccini (1974a, 1992), Hills and Brothwell (1979), Waddle (1994), Brauer and Rimbach (1990) and Stringer (1974b, 1978, 1987, 1992; 147

307, display s a columnar form frontal process and multiple (3) zygomaticofacial foramina . Vi 308 displays a large frontal sinus restricted to the well-developed supraorbital torus . The supraorbital torus is not complete enough to determine if it had the midorbital reduction identified on other Vindija frontals. An isolated upper incisor is shovelshaped. The G 1 material is "not significantly different" from the level G3 sample (Wolpoff et al. 1981:540), and fits within a Neanderthal range of morphological variation (Karavanic and Smith 1998; Smith and Ahem 1994). But, as Smith states The problem is really the paucity and fragmentary nature of the hominid remains from G 1, which renders any unequivocal interpretation of their significance impossible . ( 1984:162)

Hab good 1982, 1984a, 1984b; Smith 1976b, 1978a; Stringer etal.1984) . La Quina H5 deviates from the general Neanderthal pattern in being markedly dolichocranial (cephalic index of 67 .8), with a small cranial capacity (1,367cc) , relatively thin vault bones and a very large , and elongated occipital bun (Plate 11). Spy 2 displays variation from the general Neanderthal pattern in having a moderately high frontal squama with a steep supraglabellar segment , and relatively straight-sided parietals (Hrdlicka 1930; Thoma 1975). The Guattari-Monte Circeo 2 and 3 mandibles also deviate from the typical Neanderthal mandibular pattern (Fig. 5:16) . Guattar i-Monte Circeo 2 has single mental foramina positioned between the fourth premolar and the first molar, a small retromolar gap, and a slightly developed mental eminence , whereas Guattari-Monte Circeo 3 has multiple mental foramina positioned between the fourth premolar and the first molar, a small retromolar gap, and a slightly developed mental eminence .

Level F has also produced a small hominid sample (Karavanic and Smith 1998; Smith et al. 1985). A number of isolated anterior teeth fall within both Neanderthal and Upper Palaeolithic Homo sapiens ranges . Two parietal fragment s (Vi 204 and Vi 304) that articulate are described as displaying an even curve along the sagittal suture, a slight indication of lambdoidal flattening and in norma occipitalis a gabled contour (Karavanic and Smith 1998; Smith et al. 1985). Karavanic and Smith (1998) suggest that the parietal would not fit within the morphological range of the level G material. The identification of the level F hominid material as anatomically modem Homo sapiens would seem the most parsimonious conclusions from the limited sample .

Although there is clearly variation within the Neanderthal sample , it does present a morphologically consistent pattern that differentiates it from other hominid samples .

Vindija A large collection of fragmentary hominid remains, lithic material and a large fauna! collection have been recovered from Vindija Cave, near Zagreb, Croatia (Fig. 5:5; Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984; Smith and Ahem 1994; Smith et al. 1985; Wolpoff and Malez 1977; Wolpoff et al. 1981). Hominid material has been recovered from Mousterian and Upper Palaeolithic levels at the site. The hominid remains are very fragmentary and a detailed description of each fragment will not be undertaken here as this can be found in a number publications (Karavanic and Smith 2000; Malez et al. 1980; Smith 1982, 1984; Smith and Ahem 1994; Smith et al. 1985; Wolpoff et al. 1981). Level G3 has provided the stratigraphically earliest hominid remains from Vindija . The fragmentary hominid sample exhibits a Neanderthal morphological pattern . The fragments have well-developed supraorbital tori with large frontal sinuses restricted to the torus , ophryonic grooves, occipital tori with suprainiac fossae , bicanine expansion of the alveolus , retromolar gaps , posteriorly-angled ascending rami with rounded gonial angles , mental foramina positioned under the first molar, limited development of the mental eminence , and rounded superior orbital borders (Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984; Smith et al. 1985; Wolpoff et al. 1981). Level G 1 has also produced a number of fragmentary hominid remains (Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984; Smith and Ahem 1994; Smith et al. 1985; Wolpoff et al. 1981). The fragments of most interest include Vi 207-a right mandibular ramus, Vi 307-a left zygomatic bone and Vi 308 -a left frontal fragment. Vi 207 has a retromolar space , a horizontal-oval mandibular foramen form and a posteriorly angle ramus. The zygomatic bone Vi

Level D at Vindija contains a Gravettian assemblage and the remains of anatomically modem Homo sapiens (Malez et al. 1980; Wolpoff et al. 1981). Within the level G3 and G 1 samples there appear to be some features that vary from the typical Neanderthal pattern, although generally still within the Neanderthal range of variation. There is said to be less mid-facial prognathism (although the mandibles still display retromolar spaces and angled rami) , reduced facial size, thinner, less projecting, supraorbital tori, the beginnings of the separation of the superciliary arch from the supraorbital trigone, smaller anterior teeth (especially as regards breadth) , reduced nasal breadths , reduced alveolar heights, more vertical symphyseal regions with better-developed mental eminences, reduced nasal breadths , lack of lambdoidal flattening and occipital bunning , and possibly higher and thinner vaults (Karavanic and Smith 1998; Smith 1982, 1984; Wolpoff et al. 1981). The material is very fragmentary , and so some of these differences (such as the degree of mid-facial prognathism , occipital bunning or vault height) are only approximations based on small fragments of the relevant region. Other differences may be due to predominance of females and juveniles , which present a biased view of the population . The postcranial remains suggest small body size which may have also resulted in less pronounced expression of certain Neanderthal features. What we could have at Vindija , therefore , is a sample dominated by young females with small body sizes (see also Brauer 1989, 1992; Stringer et al. 1984). However , Wolpoff et al. (see also Smith 1984; Smith et al. 1989) stated that .. .it appears unlikely that the modem-like aspects of the Vindija G3 hominids can be 148

(Smith 1982, 1984; Wolpoff et al. 1981; Valoch 1968). It has been suggested that level G3 may represent the latter part of the "Lower Wurm stadia}" (Wolpoff et al. 1981). An amino acid racemisation date of 42,400+/-4,300 years BP has been obtained for a bone sample from level G3 (Smith et al. 1985), although there can be problems with this method when used on bone. A hominid bone from level G3 (Vi-75G3/h-203) has provided an accelerator mass spectrometry (AMS) radiocarbon date of 42,000 years BP (Ua-13873), which is consistent with the amino acid racemisation date (Krings et al. 2000).

purely attributed to allometric or other effects of size and/or sexual dimorphism. (1981:542) Ahem et al. (2002) have concluded from an analysis of supraorbital torus size, that there was a low probability that the Vindija hominids were drawn from a biased sample (i.e. a sample dominated by young and/or female individuals). Trinkaus and Smith ( 1995) have used postcranial elements (Vi 209-scapula glenoid fossa, Vi 300-3 rd/4 th proximal phalanx and Vi 203-5 th metatarsal) to investigate if the Vindija Neanderthals had a smaller body size than other Neanderthals. They concluded that although there was a suggestion that the Vindija Neanderthals were slightly smaller than other Neanderthals, this was not significant (Trinkaus and Smith 1995).

On the basis of the presence of massive-base bone points and a split-based bone point, level G 1 has been categorised as an early Upper Palaeolithic layer (Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984; Wolpoff et al. 1981). As mentioned above, the association of the bone points with the hominid remains has been questioned (d 'Errico et al. 1998; Stringer 1982; Zilhao and d'Errico 1999; but see Karavanic and Smith 1998, 2000). Level G 1 may represent a warm period and has been equated with the Podhradem ("Wiirm 1/11") interstadial (Wolpoff et al. 1981 ). Stringer (1982; comment in Smith 1982) pointed out that there are two interstadial complexes that level G 1 could be equated with. The earlier complex (Hengelo/Charbon/Podhradem) dates from 38,000 to 42,000 years BP, while the second complex (Denekamp/Grand Bois/Arey) is later and dates between 32,000 to 29,000 years BP (Stringer 1982; Woillard and Mook 1982). A radiocarbon date of 33,850+/-520 years BP (GrN-979) has been obtained from an early Aurignacian level at the Hrvatsko Zagorje site of Velika Pecina (Smith 1976a; Vogel and Waterbolk 1972), which has been used to give an approximate age to level G 1 (Smith 1982, 1984). A radiocarbon date of approximately 26,970 years BP has been obtained from a sample at the interface between Fd and Fd/d, which contains an Aurignacian assemblage (Smith et al. 1985). Level D has provided a radiocarbon date of around 27,000 years BP (Wolpoff et al. 1981).

Much debate has centred on the nature of the archaeological context of the Vindija hominids. Level G3 contains a late Mousterian industry with some tools made on blades (Karavanic and Smith 1998; Malez et al. 1980; Wolpoff et al. 1981). The limited lithic material from G 1 includes pieces typical of both the Mousterian and the Upper Palaeolithic (endscrapers, burin and a retouched blade (Karavanic and Smith 1998; but see Zilhao and d'Errico 1999). Associated with this lithic material are three massive-base bone points and a split-based bone point (Karavanic and Smith 1998). On the basis of the bone tools and especially the split-based bone point, level G 1 has been regarded, "though not conclusively", as an Aurignacian layer (Malez et al. 1980; Smith 1982, 1984; Wolpoff et al. 1981) or Szeletian (Karavanic and Smith 1998). The F complex contains what appears to be an Aurignacian assemblage (Smith et al. 1985; Wolpoff et al. 1981). The association of the G 1 hominids and the split-based bone point in particular and the other Upper Palaeolithic artefacts in general has been questioned (d'Errico et al. 1998; Stringer 1982; Zilhao and d'Errico 1999). It has been suggested that the level G 1 is quite thin and subject to postdepositional processes such as cave bear denning and cryoturbation, which could account for the mixing of material from Middle and Upper Palaeolithic layers. Karavanic and Smith ( 1998, 2000) admit that parts of the cave did experience postdepositional disturbance, but argue that there was no evidence of such processes where most of the hominid material, including Vi-207 and Vi-208, or the split-based bone point were recovered. They also contend that there is no evidence of damage on the lithic or bone tool material that would be consistent with mixing by postdepositional processes.

More recent direct dating attempts have clarified the age of the site. An AMS radiocarbon date of 33,000+/-400 years BP (ETH-12714) has been obtained from a fragment of cave bear ( Ursus spelaeus) from level G 1 (Smith et al. 1999). Direct AMS dates of 29,080+/-400 (OxA-8296) and 28,020+/-360 (OxA-8295) have also been obtained from Vi207, a right mandible fragment, and Vi-208 a left parietal fragment respectively, both from level G 1 (Smith et al. 1999). These results suggest that earlier reported dates of 51,000+/-8,000 years BP (U-Th) and 46,000+/-7,000 years BP (U-Pa) for Vi-207 were overestimates (Smith et al. 1999). The dates would place the G 1 deposits into the second interstadial complex (Denekamp/Grand Bois/ Arey) as noted above. These dates establish the Vindija hominids as some of the oldest conclusively dated Neanderthals in Europe. The dates also suggest that level GI may span at least 3,000 years.

If the identification of the archaeological assemblages were correct, Vindija would be the only site in Europe with Neanderthals associated with an Aurignacian assemblage. However, if the G 1 assemblage was a Szeletian industry, the presence of Neanderthal hominids would parallel the situation at sites such as Grotte du Renne (Arcy-sur-Cure) and Saint-Cesaire where Neanderthals have been found associated with industry combining flake, blade and bone technologies (see later discussion).

The Vindija hominid material was too fragmentary to include in any analyses. Overall, there is no feature or combination of features evident on the level G complex hominids that is inconsistent with their classification as Neanderthals. The identification of the level F hominid material is problematic due to the limit number and state of preservation, but attribution to anatomically modem Homo sapiens is a reasonable assessment at present.

Based on cultural, stratigraphical and fauna} data level G3 has been correlated with the "Lower Wiirm (Wiirm I) stadial" and dated between 40,000 and 59,000 years BP 149

evolutionary trend towards anatomically modem humans. However, there may not be a clear morphological trend towards anatomically modem humans . There is a great deal of morphological variation within the Vindija level G hominid sample as typified in the variation in the size of the retromolar space on mandibles and brow-ridge projection and thickness (Smith et al. 1985; Wolpoff et al. 1981). The hominids from Krapina seem to have lacked lamdoidal flattening and occipital bunning, which may be a feature that characterise s the Hrvatsko Zagorje Mousterian populations and not an evolutionary trend towards a more modem morphology . Also, early Upper Palaeolithic crania such as Mladec 1, 5 and 6 and even Vindija level F parietals display lamboidal flattening, and so there is no evolutionary trend in this feature as the Vindija level G material does not document the gradual developmen t of lamdoida l flattening. The Vindiji frontals are also not intermediate between earlier Neanderthals and early Upper Palaeolithic hominids in all browridge dimensions as is often suggested (Smith and Raynyard 1980 Smith et al. 1985 ; Wolpoff et al. 1981). Although early Upper Palaeolithi c crania from central Europe have supraorbita l tori with reduced thickness laterally , the male crania such as Mlade c 5 and 6 have medial segments that are very massive and are thicker than the Neanderthals , including the Vindija material (Frayer 1986; Smith et al. 1985; Wolpoff et al. 1981). The trend in reduction of the supraorbital torus identified at Vindija would be inconsistent with lateral reduction and medial segment increase in thickness, as would be required to result in the pattern evident on the Mladec male crania which Frayer (1986) demonstrates exceed the Neanderthals in medial and central dimensions. Also , if the Sala 1 frontal from the Slovak Republic is of last interglacial age (Sladek et al. 2002), lateral thinning of the supraorbital torus as identified on the Vindija material may be the continuation of a central European Neanderthal morphological trend unrelated to the origin of anatomically modem humans.

The level G hominid remain s have often been described as demonstrating the transition from Neanderthals to anatomically modem Homo sapiens (Frayer et al. 1993; Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984, 1992; Smith and Ranyard 1980; Smith et al. 1985; Wolpoff 1980a, 1992; Wolpoff et al. 1981). Smith (1992) suggested that the Vindija material was an "excellent transitional sample'', while W olpoff et al. stated ...the Vindija Neandertals vary more in the direction of modem Europeans than any other series of European Neandertal finds. (1981:541) Within the level G3 and G 1 samples there appear to be some features that deviate from the typical Neanderthal pattern , although generally still within the Neanderthal range of variation. There is said to be less mid-facial prognathism (although the mandibles still display retromolar spaces and angled rami), reduced facial size, thinner, less projecting , supraorbital tori, the beginnings of the separation of the superciliary arch from the supraorbital trigone, smaller anterior teeth (especially as regards breadth) , reduced nasal breadths , reduced alveolar heights, more vertical symphysea l regions with better-developed mental eminences , reduced nasal breadths, lack of lambdoidal flattening and occipital bunning , and possibly higher and thinner vaults (Karavanic and Smith 1998; Smith 1982, 1984, 1992; Wolpoff 1992; Wolpoff et al. 1981). However, the material is very fragmentary , and so some of these differences (such as the degree of mid-facial prognathism, occipital bunning or vault height) are only approximations based on small fragments of the relevant region. Other differences may be due to predominance of females and juveniles, which present a biased view of the population. The postcranial remains suggest small body size which may have also resulted in less pronounced expression of certain Neanderthal features. What we could have at Vindija , therefore, is a sample dominated by young females with small body sizes (see also Brauer 1989, 1992; Stringer et al. 1984). However, Wolpoff et al. (see also Smith 1984; Smith et al. 1989) stated that .. .it appears unlikely that the modem-like aspects of the Vindija G3 hominids can be purely attributed to allometric or other effects of size and/or sexual dimorphism. (1981:542) Ahem et al. (2002) have assessed the probability that the supraorbital metric difference s between the Krapina and Vindija samples are solely due to sample bias. They concluded that there was a low probability that the differences were due to sample composition bias (eg. a sample at Vindija dominated by young and/or female individuals), but rather were real populational differences. Trinkaus and Smith ( 1995) have used postcranial elements (Vi 209-scapula glenoid fossa, Vi 300-3 rd/4 th proximal phalanx and Vi 203-5 th metatarsal) to investigate if the Vindija Neanderthals had a smaller body size than other Neanderthals. They concluded that although there was a suggestion that the Vindija Neanderthals were slightly smaller than other Neanderthals , this was not significant (Trinkaus and Smith 1995).

The level G3 hominids are most probably gracile Neanderthals and do not appear to be (to quote McCown and Keith 1939) in the "throes of evolutionary change" .

Sain t-Cesaire The remains of a hominid were recovered from a Chatelperronian level at the La Roche a Pierrot rock shelter near Saint-Cesaire, Charente-Mar itime, France , which formed part of a stratigraphical sequence from Mousterian layers through to advanced Aurignacian layers (Fig. 5: 1; ApSimon 1980; Day 1986; Leveque and Vandermeersch 1980, 1981; Vandermeersch 1981b). The hominid remains consist of most of the right half of the anterior vault , face and mandible , and postcranial material (Leveque and Vandermeersch 1980, 1981; Stringer et al. 1984; Vandermeersch 1981b). Gargett (1999) has argued that the disarticulated , incomplete and fragmentary condition of the Saint-Cesaire individual suggests that it was preserved (and killed?) by rockfalls and was not an intentional burial. The cranium (Plate 12), which could be that of a female, displays most of the full suite of Neanderthal features (Table 5:6), where it can be compared . The anterior vault is low, elongated and flat, while the frontal squama is receding, and the temporal squama is low. The well-developed supraorbital region is separated from the frontal by a wide ophryonic

Wolpoff et al. ( 1981) suggest that the morphological pattern evident in the Vindija hominid sample may be related to reducing functional stress in the facial region that reflects an 150

groove and a marked depression above the glabella. The arched brow-ridges follow the contour of the orbits, and there appears to be slight lateral thinning of the right brow-ridge, although Smith ( 1984) contends that there is no brow-ridge reduction evident on Saint-Cesaire (contra Wolpoff 1992). The interorbital region is wide. The relatively gracile (compare~ to male Neanderthals) face is quite large and displays marked mid-facial prognathism. The large right orbit is more quadrilateral than is common among Neanderthals. It has an orbital index of 86.1 which makes it mesoconch. The male Neanderthals (La Chapelle 1, La Ferrassie 1 and Guattari-Monte Circeo 1) are chamaeconch, whereas the female Neanderthals (Gibraltar 1 and La Quina H5) are mesoconch (Plate 11). The right malar is inflated and lacks a canine fossa. The nasal aperture is large, although it is narrower than most other Neanderthals. It is leptorrhine (nasal index 46.6), while most other Neanderthals are platyrrhine.

Anatomically modern Upper Palaeolithic The anatomically modem Upper Palaeolithic inhabitants of Europe will not be discussed in any detail here as they are already Homo sapiens and so postdate the change from an archaic to a modem morphology. Descriptions and discussions of this material may be found in other sources (von Bonin 1935; Coon 1962; Frayer 1978, 1984, 1986; Gambier 1989; Henke and Xirotiris 1982; Hrdlicka 1930; Jelinek 1954, 1969, 1983; Keith 1925; Morant 1930; Smith 1982, 1984; Stringer et al. 1984; Wolpoff 1980a; and references therein).

Discussion As mentioned in the introduction of this chapter, Europe has been the focal point of most studies of the origin of anatomically modern humans and so many explanations have been suggested for the skeletal sample from the region. The impact of the various explanations has varied from little consideration to major debates. For example, van Yark (1983, 1984, 1985) using multivariate techniques found that Petralona, Arago 21, Steinheim and Kabwe 1 belonged to a separate lineage to that which led to modem Homo sapiens and proposed that human evolution was influenced by cosmic forces. Also, Delson (1981) has suggested that Homo sapiens could have originated in Europe. He argued that a Homo erectus group could have become isolated in Europe and that adaptation to the new environment resulted in the development of Homo sapiens (Homo sapiens heidelbergensis). These explanations have not received a great deal of support. In general, there have been four major explanations of the origin of modem humans in Europe that have received wide support.

The right half of the mandible, including most of the ascending ramus, is preserved. It has multiple mental foramina positioned under the right first molar, a welldeveloped retromolar gap, slight development of a mental eminence, and a posteriorly-angled ascending ramus with a rounded gonial angle. The postcranial material displays characteristic Neanderthal features (Leveque and Vandermeersch 1980, 1981; Stringer et al. 1984; Vandermeersch 1981b). The Saint-Cesaire hominid was recovered from the upper Chatelperronian level at the site {ApSimon 1980; Leveque and Vandermeersch 1980, 1981; Vandermeersch 1981b). Above this level were proto-Aurignacian and advanced Aurignacian layers, while below it was another Chatelperronian level and a number of Mousterian layers. The hominid is clearly from a Chatelperronian level. Radiocarbon dates for the Chatelperronian range between 30,000 years BP and 38,000 to 40,000 years BP (ApSimon 1980; Harrold 1983, 1989; Mellars 1998a, 1998b). Thermoluminescence dating has been undertaken on 20 burnt flints from the site. (Mercier et al. 1991). Flints from the layer containing the hominid remains provided dates ranging from33,700+/-5,400 years BP to 38,200+/-5,300 years BP with an averaged date of 36,300+/-2,700 years BP (Mercier et al. 1991).

The Pre-sapiens Hypothesis The Pre-sapiens Hypothesis was initially proposed by Boule (1913) who thought that there was a morphological and cultural replacement in Europe of the Neanderthals by anatomically modem Homo sapiens with Upper Palaeolithic assemblages. Keith (1915, 1925) also saw the "classic" Neanderthals as a side branch that became extinct. The Presapiens Hypothesis was more fully developed by Vallois (1949, 1954, 1958; Boule and Vallois 1957) who identified two independent lineages in Europe. The pre-Neanderthal line included Mauer, Steinheim, Montmaurin, Ehringsdorf and Saccopastore, while the pre-sapiens line consisted of Swanscombe and Fontechevade. Vallois stated It is scarcely to be doubted that the Presapiens provided the origin of sapiens man of the European Upper Palaeolithic. (1958:494) The pre-sapiens and anatomically modem Homo sapiens are said to have evolved "Somewhere in the east, doubtless in Western Asia" (Vallois 1958:495). Swanscombe and Fontechevade are identified as either "emissaries" of an Asiatic stock or evidence of a local European lineage that developed in parallel with the pre-Neanderthals. In both cases the pre-sapiens finally gave way to he Neanderthals. Vallois contended that

The Saint-Cesaire partial cranium linked, initially, with La Ferrassie 1, and then with the other Neanderthal crania. The mandible, however, was slightly removed from the major Neanderthal group (Fig. 5:16). This separation is caused by the slight chin development of the Saint-Cesaire mandible. Saint-Cesaire fits within the Neanderthal range of morphological variation and displays most of the typical Neanderthal characteristics where it can be compared. It has been suggested that Saint-Cesaire resembles the Vindija and Sala specimens from central Europe (Smith and Ranyard 1980; Wolpoff 1981; Wolpoff et al. 1981), and so is transitional, or demonstrates the evolutionary trends towards anatomically modem humans within the Neanderthals (see Wolpoff and Frayer 1992). However, in most respects SaintCesaire demonstrates little change in morphology from the earlier "Wiirm" Neanderthals and should not be regarded as transitional. 151

anatomically modem Pleistocene .

...the Presapiens lost no time in taking a fmal revenge on their Mousterian conquerors. (1958:495) This explanation can no longer be substantiated in that the morphology of the only two pre-sapiens can be seen ~o be Neanderthal in overall configuration (Brauer 1984c; Strmger 1974b; Stringer et al. 1984; Wolpoff 1980a). Vallois' reconstruction of the Fontechevade hominids without browridges has been seriously questioned (discussed above; Brace 1964; Corruccini 1974a; Howell 1957; Sergi 1953; Trinkaus 1973). Swanscombe displays a characteristic Neanderthal configuration of the occipital torus (Santa Luca 1978). It has also been demonstrated that there are many similarities between Swanscombe, a pre-sapient, and Steinheim, a preNeanderthal (Breitinger 1964; Habgood 1982, 1984a, 1984b; Habgood and Walker 1986; Howell 1960; Mora~t 1938; Stringer 1974b, 1978; Weidenreich 1943a; Werner and Campbell 1964; Wolpoff 1980a, 1980b). There is, therefore , no pre-sapiens upon which to base this explanation.

The Spectrum Hypothesis The Spectrum Hypothesis was developed by Weiner (1958 ; Weiner and Campbell 1964), but has been supported by others (Cuenca and Walker 1980, 1981; Habgood and Walker 1986; Stringer 1974b). This explanation argues for a morphological continuity through time and space, and proposes that following the Homo erectus stage there was present a spectrum of varieties of early Homo (?re Rhodesian, pre-Solo, pre-Neanderthal and pre-sap1ens forms). The pre-Neanderthal forms included the mandibles from Mauer and Montmaurin, whereas Swanscombe and Steinheim were regarded as pre-sapiens. Some of the forms would have continually intergraded while others would have been subjected to varying degrees of isolation. It contends that by the Upper Pleistocene the spectrum had broadened with distinctive populations of the Solo type, Rhodesian type, Neanderthals , "intermediate" forms such as Skhiil and Krapina, and Homo sapiens proper ("Cro-Magnon man") . Weiner and Campbell state The last alone [Cro-Magnon man] survived as a single species, to give rise to the modem racial and regional varieties . (1964:206) There are problems with this explanation. The similarities between Swanscombe , Steinheim and the Neanderthals have been discussed above. The Skhiil and Krapina sample do not appear to be "intermediate" forms. The former are clearly anatomically modem Homo sapiens with a few archaic features, whereas the latter are defmitely Neanderthal in overall morphological configuration. This hypothesis has recently supported in that it has been argued that the Atapuerca-Gran Dolina hominids represent the last common ancestor of Neanderthals and modern humans in Europe and have been assigned to the new species - Homo antecessor (Arsuaga et al. 1999; Arsuaga et al. 2001; Bermudez de Castro et al. 1997). However , there are similarities between the Atapuerca-GD hominids and other Middle Pleistocene hominids in Europe and , as outline within this chapter, there is no evidence to support a contention that there were two morphologically distinct lineages evolving within Europe prior to the appearance of both "classic" Neanderthal and

Homo

sapiens during the Late

Also, anatomically modern Homo sapiens do not appear to have evolved in Europe (discussed below).

The Replacement Hypothesis Multiregional Hypothesis

and

the

Both of the two remaining explanations of the origin of anatomically modem humans in Europe identify in situ evolution of the Neanderthals. They differ in that one explanation suggests a mid Upper Pleistocene replacement of the Neanderthals, whereas the other contends that there was a local transition into anatomically modem Homo sapiens . These two explanations are still hotl y debated.

The Replacement Hypothesis The Replacement Hypothesis (formerly know as the PreNeanderthal or Rapid Replacemen t Hypothesis, but now popularly know as the Out of Africa Hypothesis) contends that Homo sapiens evolved outside Europe and towards the middle of the Upper Pleistocene moved into Europe and replaced the Neanderthals . The Neanderthals were thought to have played little, if any part, in the origin of anatomically modem humans. The place of origin of the modem humans is placed in western Asia and/or southern Africa. Sergi (1953) highlighted the polytypic nature of the preNeanderthals (in contrast to the relatively homogeneous "classic" Neanderthals) and suggested that both the Neanderthals and anatomically modern humans developed out of this earlier more "generalised" group. Howell (1951, 1952, 1957) also identified early "generalised" Neanderthals and suggested that a group of this type became isolated in western Europe during the last glacial and developed into the "classic " Neanderthals. He states Further to the east, from the Mount Carmel population and similar groups, modern man evolved . (1951 :412) Le Gros Clark , who also supported this explanation, stated ...H. neanderthal ensis arose as an aberrant (and, in some respects, a retrogressive) collateral line from a pre-Mousterian or early Mousterian variety of H. sapiens ; and...the former species eventually became extinct. (1964:76) Howells (1970 , 1974, 1975, 1976a, 1981) has supported this view with both morphological and metrical studies. After perfonning a multiple discriminant analysis Howells concluded that the Neanderthals were ...statistically speaking , unusual or impossible members of modern man ...[and that]...The Upper Paleolithic and later Europeans see~ more clearly distinguished than ever from theu Neanderthal predecessors . (1970:271) Subsequently, Howells stated that 152

...for much of the Wiirm the two distinct populations existed in different places, until the Neanderthals were replaced in Europe by H. sapiens sapiens ... (1981 :76)

Stringer (1974b, 1978, 1982, 1984b, 1985, 1989a, 1989b; Stringer et al. 1984; Stringer et al. 1981) has been a strong advocate of the Replacement Hypothesis. He argues there is a major morphological discontinuity in western Europe which he emphasises in his arguments. After a major multivariate study he stated ...any attempt to derive Upper Palaeolithic and recent man from the later Neanderthals ...would involve some remarkable evolutionary twists and turns in cranial, not to mention postcranial evolution. (1974b:333) Stringer also carried out a morphological-cladistic study and concluded that anatomically modem Homo sapiens did not originate in Europe and that No genuinely transitional fossils displaying combinations of Neanderthal and modem derived characters have been found in Europe or southwest Asia ... (1985:294)

Boaz et al. ( 1982) have argued that anatomically modem humans were forced into Europe by increasing aridity in north Africa and western Asia during the Upper Pleistocene, and that they swamped-replaced the small and geographically isolated population of Neanderthals (but see Fogarty and Smith 1987). Turner (1985, 1987) has also supported the Replacement Hypothesis. His examination of the teeth of some Neanderthal and Upper Palaeolithic anatomically modem humans did not identify evidence of continuity in Europe (Turner 1985). Turner also states My own preliminary dental morphological studies of Old World populations suggest that Southeast Asia could have been the geographical node for all branches of modem populations ... (Turner 1985:5)

Stringer et al. express a similar view stating that western Europe has no good evidence for the actual origin of a.m. H sapiens. (1984:115)

The Replacement Hypothesis has . also been strongly supported by Vandermeersch (1981a, 1981b, 1981c, 1982; Stringer et al. 1984), Brauer (1980a, 1980b, 1981, 1982, 1984a, 1984b, 1984c, 1989) and Stringer (1974b, 1978, 1982; 1984b, 1989a, 1989b; Stringer et al. 1984; Stringer et al. 1981).

They conclude that Given the morphology and dating of the known material, it is still possible that the modem characters of the late Upper Pleistocene hominids of this area [western Europe] were ultimately derived from an exotic source such as Africa. (1984:121)

Vandermeersch (1982; Bar Yosef and Vandermeersch 1981; cf. Chapter 4) has argued (even prior to the revised dating of many of the western Asian sites; cf. Chapter 4) that the anatomically modem Jebel Qafzeh hominids are earlier than most of the Neanderthals and so anatomically modem humans cannot be the descendents of the Neanderthals. He also contends that there were two separate lineages in western Asia. One, consisting of Zuttiyeh, Jebel Qafzeh and Skhiil, is Homo sapiens, while the other, made up of the material from Tabiin, Amud and Shanidar, is Neanderthal. Vandermeersch states Fewer morphological transformations are required to proceed from the evolved Homo erectus to Homo sapiens than to proceed from Neanderthal to Homo sapiens sapiens ... [and that] ... The Neanderthals represent a divergent and very specialized branch in our history. (1982:297)

The Neanderthal Hypothesis

Phase

or

Multiregional

The Neanderthal Phase of Man, Regional Continuity or Multiregional Hypothesis was redeveloped from an earlier theory by Brace ( 1962, 1964, 1966, 1979), who stated ...I suggest that it was the fate of the Neanderthal to give rise to modem man ... (1964:19) Schwalbe, who is attributed with initially proposing this hypothesis, arranged the fossil hominids available to him as stages in a linear sequence which he believed represented the course of human evolution (Smith 1987). He placed the Neanderthals in an intermediate position between modem humans and Homo erectus. He suggested In a pure zoological sense Homo primigenius [the Neanderthals] is an intermediate between Homo sapiens and Pithecanthropus erectus. (1906: 14; quoted in Smith 1987:3) and that ...if Homo sapiens appears later than Homo primigenius [the Neanderthals], the more probable perspective is the development of the

Brauer ( 1984a, 1984b, 1984c, 1989) has proposed what he calls the Afro-European sapiens Hypothesis. This explanation contends that anatomically modem humans originated in sub-Saharan Africa and subsequently moved north during the last glacial and replaced the Neanderthal populations in north Africa and western Asia ( cf. Chapters 2, 3 and 4). Brauer suggests that During the next millennia - in what was probably a relatively slow process of hybridization and replacement - they also superseded the European Neandertals. ( 1984b: 158)

former from the latter, or an unknown form similar to it... (1923:297; quoted in Smith 1987:5) 153

clearly documented information ...

Hrdlicka (1927, 1930; Spencer 1984; Spencer and Smith 1981) supported Schwalbe's unilineal sequence and stated that there was ...the evolution of the Neanderthaler into later man ...[and that]. ..there appears to be less justification for the conception of a Neanderthal species than there is for that of a Neanderthal phase of Man. (1930:348)

by

the

available

(1984:192-3) and with Ranyard he suggested the ...data presently available are best interpreted to indicate that Homo sapiens neanderthalensis evolved directly into Homo sapiens sapiens at least in South-Central Europe. (1980:607-8)

Weidenreich (1940 , 1943a, 1943b, 1947) also placed the available hominid sample within a unilinear evolutionary sequence identifying a world-wide Neanderthal stage through which hominids had to pass so as to become anatomically modem. He stated There is now evidence that: ( 1) Neanderthaloids have been directly transformed into Homo sapiens forms; (2) Neanderthalians should not be considered to have become extinct without leaving any descendents behind. (l 943b:277) Weidenreich (1947:201) produced a diagram to illustrate his views. Missing from this diagram, however , are the "classic,, Neanderthals . Instead Weidenreich has an evolutionary sequence progressing from Tabfin through the Skhul group to Upper Palaeolithic European hominids.

The Neanderthals: Where From and Why? During the first half of last century there was a major debate over the "fate of the Neanderthals" (Hammond 1980, 1982; Spencer 1984; Spencer and Smith 1981; Smith 1987). This situation has not changed even though our knowledge and understanding of this period in human evolution has greatly increased . At this point it is instructive to quote Howells who observed that The Neanderthal problem is as much a problem of anthropologists as of Neanderthals. ( 1973a: 103) Stringer has even suggested that If we cannot solve the "Neanderthal Problem " and thereby arrive at an understanding of the relationship of Neanderthal to "modem,, Homo sapiens ...there would seem little hope of resolving any of the more complex issues concerning hominid evolution . (1982:431)

Coon (1962) supported the arguments of Weidenreich and Hrdlicka and argued for the contribution of the Neanderthals to the origin of modem European populations. He stated The Neanderthals became extinct; of that there is no question. But their extinction was probably of the usual human form, extinction by absorption. (1962:549)

Neanderthal Origins Which of the two hypotheses - Replacement or Regional Continuity - best explains the European hominid record? The first question that needs to be addressed is the origin of the Neanderthals. All of the evidence indicates that the Neanderthals evolved in Europe with the gradual appearance of the characteristic Neanderthal morphological configuration (Fig. 5:6; Table 5:6; Hublin 1984; de Lumley, M-A. 1973; Smith 1985; Stringer 1985, 1989b; Stringer et al. 1984; Vandermeersch 1985; Wolpoff 1980a, 1980b, 1989a).

Eastern European scholars have tended to support an in situ origin of anatomically modem humans (GorjanovicKramberger 1906; Jelinek 1969, 1985; Radovcic 1985). For example, Jelinek stated ...the appearance of H. sapiens sapiens in Central and Eastern Europe ...need not be explained in terms of a sudden migration from East to West, but rather in terms of local evolution in populations ... (1969:492)

The "primitive" European morphological pattern is revealed by the mid Middle Pleistocene hominid sample (AtapuercaGD, Mauer , Petralona, V ertesszollos, Arago, B ilzingsle ben[?]) .

The Multiregional Hypothesis has been supported by a number of authors (cf. Spencer 1984; Spencer and Smith 1981), but at the forefront of this group have been W olpoff (Brose and Wolpoff 1971; Wolpoff 1971b, 1979, 1980a, 1981, 1989a, 1989b; Wolpoff and Malez 1977; Wolpoff et al. 1981), Frayer (Frayer 1986, 1992; Frayer et al. 193, 1994) and Smith (Spencer and Smith 1981; Smith 1978a, 1978b, 1982, 1983, 1984, 1985; Smith and Paquette 1989; Smith and Ranyard 1980; Smith et al.1989; Trinkaus and Smith 1985). All three highlight what they see as evolutionary development from late Neanderthals to early anatomically modem humans especially in central Europe.

The Atapuerca-GD hominids display archaic features reminiscent of other Middle Pleistocene hominids from Europe and Africa and do not have any of the characteristic Neanderthal features. The Petralona cranium displays predominantly "primitive" features and resembles archaic hominids from other regions such as sub-Saharan Africa (Bodo 1 and Kabwe 1). It does have some features of the splanchnocranium that become characteristic of the Neanderthals in that it displays expanded maxillary sinuses, lack of canine fossae , incipient mid-facial prognathism and a lack of angulation at the junction of the malars and maxillae. The neurocranium does not display the characteristic Neanderthal pattern , especially evident in the occipitomastoid region.

For example, Smith stated In my opinion , morphological continuity between Neandertals and the EMH [early modem human] sample in central Europe is 154

Arago 21 displays a similar pattern to that of Petralona, although it does have a sharp lower nasal border reminiscent of the Neanderthals. The Mauer mandible does not display any characteristic Neanderthal features. Apart from the posterior positioning of its mental foramen, Arago 13 does not display Neanderthal features. It is more similar to the Mauer mandible than to the "classic" Neanderthals. Arago 2, does display a retromolar gap and mental foramina positioned under the first molars. This configuration suggests that the cranium that Arago 2 articulated with, had mid-facial prognathism.

tripartite occipital torus and suprainiac fossa on La ChaiseSuard 2. The Reilingen calvaria bas maximum breadth at the supramastoid crests presenting a pentagonal shape as opposed to an "en bombe" contour, but displays an angled occipital squama presenting an incipient occipital bun and a developed occipital torus with a well-defined suprainiac fossa. This late Middle Pleistocene hominid sample may be regarded as pre- or anteneanderthals in that they do, to varying degrees, have features that become typical of the later Neanderthals. The earlier hominids (Steinheim and Swanscombe) generally do not display these Neanderthal features to the same extent as the later remains from Biache and La Chaise-Suard. Other material from this period also displays Neanderthal features. For example the stratified upper molars from Pontnewydd Cave, Wales, display taurodontism (Stringer 1984c). Although this hominid sample displays Neanderthal features, it is still differentiated from later Neanderthals in the analyses of metrical and nonmetrical data (where included). Steinheim, Swanscombe and La Chaise-Suard 2 (the only members of this group analysed) were separated from the Neanderthals, and fell with the earlier hominids such as Petralona and V ertesszollos 2 (cf. Figs. 5:7, 5:8, 5: 14, 5: 15, Table 5:3).

Vertesszollos 2 does not have the typical Neanderthal occipitomastoid configuration. Hublin (1984) has suggested that Vertesszollos 2 represents the primitive occipital pattern from which the later Neanderthal configuration could have developed. The hominid material from Bilzingsleben does not display Neanderthal features. None of these hominids could be mistaken for a Neanderthal (after a detailed examination at least; cf. Poulianos 1967). In the multivariate analyses this group (where included) was clearly separated from the "classic" Neanderthals. This separation was evident in the analyses of both metrical and non-metrical data (cf. Figs. 5:7, 5:8, 5: 14, 5: 16, Table 5:3). Neanderthal features (Table 5:6) become more evident on the late Middle Pleistocene hominid sample from Europe.

The last interglacial or early last glacial hominid sample from Europe (K.rapina, Saccopastore and Gibraltar 1) is essentially Neanderthal in overall morphological configuration. This is clearly demonstrated in the analyses of metrical and non-metrical data (Figs. 5:7, 5:8, 5:9, 5: 10, 5:11, 5:12, 5:13, 5:14, 5:15, Table 5:3). However, this material is, in some respects, more "generalised", "progressive" or less "specialised" than the later "classic" Neanderthals (Habgood 1984a, 1985a; Habgood and Walker 1986; Howell 1951; Sergi 1958). These differences are highlighted by the analyses of metrical data where, on occasions, this material is separated from the "classic" Neanderthals and placed with the earlier European hominids (Figs. 2:5, 2:6, 2:7, 2:8, 4:9, 5: 14, Table 5:3; Habgood 1982, 1984a, 1984b, 1985a; Habgood and Walker 1986).

Steinheim presents a Neanderthal occipitomastoid region and a non-Neanderthal splanchnocranium. The preserved splanchnocranium has sub-rectangular orbits, a canine fossa on the preserved right malar, a moderate amount of angulation between the malar and maxilla and only limited mid-facial prognathism. The occipital, however, has a tripartite occipital torus with suprainiac fossa. Swanscombe displays a Neanderthal occipital pattern with a tripartite transverse torus and a suprainiac fossa above the flattened medial segment. It also has a high degree of prelambdoidal flattening, which would appear to be the beginning of the development of the Neanderthal occipital "chignon". In norma occipitalis, Swanscombe also presents a rounded "en bombe" transverse contour. The Atapuerca-SH hominid sample presents a mosaic of archaic and Neanderthal features. Some of the mandibles have mental foramen under M 1 and retromolar spaces. The crania have occipital tori with suprainiac fossa, but have maximum breadth low down on the vault and parallel vault walls, not the more rounded Neanderthal configuration and do not display lambdoidal flattening or occipital buns. A well-developed occipital bun is evident on the hominid from Biache. This specimen also has a tripartite occipital torus with a suprainiac fossa and large occipitomastoid crests. In norma occipitalis Biache also displays a rounded "en bombe" transverse contour. The hominid sample from La Chaise-Suard displays a predominance of Neanderthal morphological features such as the "en bombe" contour of La Chaise-Suard 1 and the

The chronological placement of the Fontechevade and Ehringsdorf hominids is problematical. They could belong to the late Middle Pleistocene or to the early Upper Pleistocene. Their dating does not influence the morphological pattern of the hominids, which, apart from some of the parietals from Ehringsdorf, can be easily matched within the Neanderthal sample. During the first half of the last glacial (Table 5: 1; "Wurm" through "Wurm 11/111")the Neanderthal morphological configuration became established in Europe (Hublin 1984; Smith 1982, 1984; Stringer 1982; Stringer et al. 1984; Wolpoff 1980a). The Regourdou mandible, the only definitely "Wurm I" Neanderthal analysed (although K.rapina, Saccopastore and Gibraltar 1 may be from this period also), displays the characteristic Neanderthal retromolar gap, mental foramina positioned under the first molars and a receding symphyseal region with little development of a mental eminence. It was found to group with other Neanderthal mandibles in the analysis of mandibular non-metric traits (Fig.5: 16). The later "Wurm II" 155

glacial environment. The short distal limb segments of the Neanderthals would have reduced the surface area of the Neanderthal body and so lessened heat loss (Table 4:8; Trinkaus 1981).

Neanderthal sample forms a group that is separated from both earlier and later hominids on analyses of cranial metric and non-metric, and mandibular non-metric data (cf. Figs. 5:7, 5:8, 5:9, 5:10, 5:11, 5:12, 5:13, 5:14, 5:15, 5:16, Table 5:3). The latest known western European Neanderthal, SaintCesaire, also displays a Neanderthal morphological configuration (Fig. 5: 16).

Experimental studies of modern cranial form and the postcrania of rats (Steegmann 1965, 1970, 1972; Steegmann and Platner 1968) have been used to support the cold adaptation explanation of the Neanderthal morphology (but see Smith 1983; Smith and Paquette 1989). However, Neanderthal body proportions are probably the only features that can be easily attributed to climatic adaptation.

As has been discussed, although there is variation within the European Neanderthal sample, it displays a characteristic morphological pattern (Table 5:6) that differentiates it from the hominids that precede and postdate it, and from its contemporaries from Africa and Asia, although the western Asian Neanderthals are similar (cf. Chapter 4).

Hublin ( 1998) has proposed a variation to the cold adaptation model by stressing the genetic impact of population fluctuations caused by extreme climatic fluctuations ( founder effect, genetic drift and fixation of derived features). Hublin ( 1998) proposed that a cold glacial peak during oxygen isotope stage 12 (Table 5) may have resulted in significant population reductions in Europe that could have triggered the evolutionary divergence of European hominids that ultimately led to the Neanderthal morphological configuration.

As can been seen from this discussion, there is a chronological and morphological sequence that documents the gradual (over at least 200,000 years) development of the European Neanderthals. The origin of the Neanderthals is therefore, clearly documented within Europe. There is only evidence of a Neanderthal lineage in Europe during this period and so there are no "pre-sapiens" . There also does not appear to be a "spectrum" of forms in that after the late Middle Pleistocene all of the hominid remains (except Alicante if it actually had a short splanchnocranium and was an Upper Pleistocene fossil) can be placed in a Neanderthal lineage. The Mauer mandible and Vertesszollos 2 appear to display the "primitive" European morphological pattern from which it is possible to derive the Neanderthal pattern (Hublin 1984; Stringer 1985; Vandermeersch 1985; Wolpoff 1980a). Petralona and the Arago material display a combination of both Neanderthal features and characteristics that are probably the "primitive" condition for European hominids. The Bilzingsleben remains, due to their fragmentary condition, are difficult to place in this sequence in that they display archaic morphological features. However, as Stringer ( 1980, 1981) has demonstrated, the relevant parts of Petralona are similar to the Bilzingsleben remains. Neanderthal Morphology Various explanations have been put forward to explain the development of the unique Neanderthal morphological pattern, especially that of the cranium. Howell (1951, 1952) postulated that the Neanderthals may have been a product of adaptation to a very cold environment. This idea was developed by Coon ( 1962) who listed features that he thought demonstrated this adaptation to a cold environment. These features included large infraorbital foramina so as to supply additional blood to the face, and a mid-facial complex (mid-facial prognathism, expanded maxillary sinuses and large nasal aperture) that moved the nasal cavity away from the arteries that feed blood to the brain and would warm and humidify the dry cold inhaled air. Brose and W olpoff ( 1971) added large frontal sinuses, occipital bunning and body build (short and heavily built) to the list of features of cold adaptation. They state The resulting total morphological pattern for the "classic" group is one of cold adaptation built upon the morphology of the Neandertal grade of evolutionary development. (1971:1178)

The other major explanation for the development of the unique cranial morphology of the "classic" Neanderthals is functionally based. Brace (1962, 1964) proposed that the Neanderthals used their large anterior teeth for para- and non-masticatory functions and that the unique facial morphology reflects the extra force placed on the anterior dentition. The large anterior dentition with a high frequency of shovel-shaped incisors, basal tubercles, microflaking, enamel chipping, labial rounding wear, a greater degree of attrition of the anterior as compared to the posterior dentition and osteoarthritis of the temporomandibular joints are used to indicate massive anterior dental loading and a nonmasticatory use of the anterior teeth. This suggestion has been widely supported (Brose and Wolpoff 1971; Mann and Trinkaus 1973; Rak 1986; Smith 1983; Smith and Paquette 1989; Smith et al. 1989; Trinkaus 1983b, 1986; Wolpoff 1980a), but has also been criticised (Puech 1981; Smith, P. 1976, 1977a, 1977b; Wallace 1975). A large and heavily buttressed face (alveolar buttress, the thickened lateral orbital margin, large supraorbital torus) would be necessary to accommodate the large anterior teeth and to dissipate the vertical occlusive forces produced during the use of the anterior dentition as a tool (Brace 1962, 1964; Brose and Wolpoff 1971; Endo 1966, 1970; Heim 1976, 1978; Smith 1983; Smith and Paquette 1989; Smith et al. 1989; Wolpoff 1980a). Occipital bunning (producing a flat nuchal plane) and the unique Neanderthal occipitomastoid configuration may reflect increased musculature of the posterior cranium so as to compensate for the forces produced during anterior dental loading (Smith 1983; Smith and Paquette 1989). There is, however, some debate over the mechanism(s) that caused the formation of the Neanderthal facial form. Trinkaus (1983a, 1984a) and Smith (Smith and Paquette 1989) contend that Neanderthal mid-facial prognathism was the result of a posterior migration of the masticatory muscles and zygomatics (the lateral face) in relation to the midline, without any decrease in total facial prognathism. Rak ( 1986) on the other hand, sees a change in the orientation of the

Trinkaus (1981 :218) has demonstrated that Neanderthal limb proportions "may indicate a long-term adaptation" to a 156

infraorbital region from the coronal plane to the sagittal lane. As indicated above, all see the changes as due to anterior dental loading.

reduction in various aspects of the cranium. The frontal squama is said to be higher in the late Neanderthals. A general reduction in the dimensions of the supraorbital torus is identified. This reduction is especially evident in the midorbital segment, which Smith regards as indicating an incipient division of the torus into a supraorbital trigone and a superciliary arch. Smith also identifies a reduction in facial size, robusticity and degree of mid-facial prognathism. The anterior dental dimensions also reduce. Smith (1983; Smith and Paquette 1989) explains this pattern of reduction by postulating a change in functional/adaptive demands placed on the Neanderthal face that resulted from a change in behavioural patterns. This essentially involved a decrease in the use of the anterior dentition for non-masticatory functions. This change in behaviour is said to have been caused by technological advances. It is worth noting that Smith et al. ( 1989) identified a different pattern in browridge reduction in western Europe and central Europe. In central Europe they found gradual change through time of the supraorbital torus, which suggested no abrupt change from Neanderthals to anatomically modem humans. However, in western Europe there was evidence of an abrupt change in some features of the supraorbital torus with the appearance of anatomically modern humans. Smith et al. ( 1989) concluded that, with respect to the form of the supraorbital torus, there was no evidence of replacement in central Europe, but that replacement of the Neanderthals by anatomically modern humans may have been a factor in western Europe.

The origin of the unique Neanderthal morphological configuration is not of direct importance in a discussion of the origin of anatomically modern humans. Suffice it to say that the Neanderthal pattern is unique and its development probably involved both cold adaptation and biomechanical requirements such as anterior dental loading as well as other genetic processes. The Transition (?) There can be no doubting that the Neanderthals developed slowly in Europe. The major problem is ascertaining what happened to them. They were either replaced by anatomically modern humans from some other region, evolved in situ into anatomically modern humans or some combination of continuity and replacement.

Wolpoff (1980a), Smith (1984, 1983; Smith and Paquette 1989; Smith and Ranyard 1980) and Trinkaus ( 1983b, 1984a, 1986, 1989; Trinkaus and Smith 1985) have proposed a series of biomechanical-functional changes that document the change or transition from a Neanderthal morphological pattern to an anatomically modern human pattern. Wolpoff (1980a) contends that technological improvements during the Middle Palaeolithic were responsible for a trend in reduction in general body strength as reflected in changes in limb form, and reductions in bone thickness, buttressing, muscle attachment areas and overall robustness. He also suggested that the technological improvements allowed a decrease in the use of the anterior dentition for nonmasticatory functions, which instigated a reduction in the size of the anterior dentition and the supporting structures of the face (for example, the maxillary region, the supraorbital region, and the degree of mid-facial prognathism). The final consequence was a redistribution of cranial mass from the low elongated Neanderthal form to the shorter and higher form that typifies anatomically modern crania. W olpoff (1980a) identified these trends within the earlier Neanderthal sample, and so the changes were occurring before and after "the transition", not as a result of a "transition".

Trinkaus ( 1983b, 1984a, 1986, 1989) has provided the most detailed account of the reasons for a change from the Neanderthals morphology to that of anatomically modern humans. As with Wolpoff and Smith, Trinkaus contends that a lessening of anterior dental loading caused the changes in the splanchnocranium. He also details changes in postcranial anatomy, which are fundamentally related to a decrease in muscular hypertrophy. In the upper limb there was a decrease in habitual strength levels and a change in deployment and grip. In the lower limbs there was a decrease in muscular hypotrophy and an increase in the length of the distal limbs, which involved a shift away from strength and endurance to more efficient locomotor pattern. Trinkaus (1983b, 1984a, 1986, 1989) also contends that cultural improvements instigated these anatomical changes.

Smith (1983, 1984; Smith and Paquette 1989; Smith and Ranyard 1980; Smith et al. 1989; Trinkaus and Smith 1985) has mainly concentrated on the hominid sample from central Europe. He perceives a consistent pattern of change in a number of cranial features from early Neanderthals to late Neanderthals that are in the direction of anatomically modem humans. It is also of interest to note that following a detailed analysis, Smith (1976b, 1978a) was unable to identify any evolutionary trends within the Krapina sample, which may span 30-50,000 years (although Krapina A is said to approach early modern humans in some features cf. Coon 1962; Minugh 1983; Minugh-Purvis and Radovcic 1991; Rink et al. 1995; Wolpoff 1980a and discussions on the proposed transitional nature of Krapina A). Therefore, if there were evolutionary trends towards an anatomically modem human morphology, the trends must have occurred after the date of the Krapina material.

There is also the possibility that the trend toward morphological reduction in the splanchnocranium, that is said to cross the Neanderthal-anatomically modem human boundary, is false. If one considers a reduction in supraorbital torus development, there is the possibility that a group of anatomically modem humans with relatively small brow-ridges (compared to Neanderthals) moved into central Europe, where there were Neanderthals undergoing a reduction in the development of the supraorbital tori. If one placed these hominids in a unilinear sequence, there could be the false impression of a trend for brow-ridge reduction across a non-existent transition. Such a scenario could also be applicable to dental reduction across the transition (compare Frayer 1978 and Wolpoff 1989a with Stringer 1982, 1989a, 1989b). Changes within the Neanderthal morphological pattern at different rates at different places throughout Europe, is most probably a continuation of morphological change and

Smith (1982, 1983, 1984; Smith and Paquette 1989; Smith and Ranyard 1980; Smith et al. 1989) does document a 157

Neanderthal features such as the squarish shape of the right orbit, slight thinning of the right brow-ridge and a slightlydeveloped mental eminence, but these can be matched on other Neanderthal crania .

gracilisation that had been occurring throughout the World during the Middle Pleistocene and not related to the origin of anatomically modem humans. Those authors who argue that in situ evolution from Neanderthals to anatomically modern humans occurred in Europe have identified supposedly transitional specimens (transition is taken to mean a morphology that is evolving from the Neanderthal type to the anatomically modem form). Arguments for a transition have focused on either identifying "transitional Neanderthals " or the persistence of Neanderthal features on early anatomically modern hominids. Hrdlicka ( 1930) identified the two hominids from the Belgium site of Spy as transitional specimens stating No better demonstration could have been furnished ...of the transitional potentialities among the later Neanderthal representatives , to which the skeleton s evidently .belong , toward s the modem human type. (1930:202) Thoma (1975), however, has demonstrat ed that the Spy hominids are not transitional specimens , but may be placed within the Neanderthal range of morphological variation. Multivariate analyses also demonstrate the Neanderthal nature of the Spy material (Figs. 5:15, 5:16; Habgood 1982, 1984b). Frayer et al. have argued that Evidence represented in the fossil record of late Pleistocene Europe demonstrates a persistence of so-called unique Neanderthal features over time, bridging the populations of the Mousterian and Upper Paleolithic. (1993:29) Wolpoff (1980a, 1989a, 1992; Frayer et al. 1993) has identified a number of what he regards as transitional specimens. As previously discussed, Wolpoff (1980a, 1992; see also Minugh-Purvis and Radovcic 1991) contends that the juvenile Krapina A is transitional because it displays only slight lambdoidal flattening on the parietals and differentiated medial and lateral elements on the supraorbital region. However, Smith (1976b) felt that morphological and metrical comparisons indicated that Krapina A was an immature Neanderthal (but see Rink et al. 1995) and Minugh-Purvis (1998) did not identify close morphological similarities between Krapina A and Skhul 1. Wolpoff (1980a, 1989a; Frayer et al. 1993) also identified as morphologically transitional Saint-Cesaire, the remains from Vindija, the ulna, maxilla , and the dental material from Grotte du Renne {Arcy-sur-Cure), Sipka and Hortus. Apart from Saint-Cesaire, this material is very fragmentary and so offers only limited morphological information . Frayer et al. (1993:29) suggested that Saint-Cesaire "shows a number of features approaching the 'modem ' condition ". However, the Saint-Cesaire partial cranium and mandible were found to group with other European Neanderthals on the multivariate analyses . This placement is consistent with the cranial and postcranial morphology of the specimen (Leveque and Vandermeersch 1980, 1981; Stringer et al. 1984). Saint-Cesaire does show a few "non-typical"

The fragmentary Vindija hominid remains have been discussed earlier and so it is not necessary to repeat the details again. In general, the material is thought to demonstrate reduced facial size, mid-facial prognathism, and brow-ridge development , the beginnings of the separation of the superciliary arch from the supraorbital trigone, slight chin development, smaller anterior teeth, higher frontals and a lack of occipital bunning (Frayer et al. 1993; Smith 1982, 1984, 1992; Smith and Ranyard 1980; Smith et al. 1985; Wolpoff et al. 1981). The material is very fragmentary and so some of these observation s are based on very small portions of the relevant morphological component. There also remain s the possibility that there is a predominance of smallish female or young individuals at the site, which could provide a biased representation of the overall morphological pattern (but see Ahem et al. 2002; Trinkaus and Smith 1995; Smith 1984; Smith et al. 1989). It is also not surprising that there is a lack of occipital bunning at this site because the preceding Neanderthals from the region (Krapina) also lack occipital buns (Smith 1976b, 1978a, 1982, 1984). A reduction in anterior dental loading could also produce a reduction in anterior dental size, brow-ridge size and degree of mid-facial prognathism (Brace 1979; Russell 1985; Smith 1983; Trinkaus 1986). This process of reduction would have been assisted if the population had higher frontal squamae (Endo 1966, 1970; Russell 1985; Smith 1983; Trinkaus 1986). Many of these "transitional" features are also consistent with evolutionary trends that Wolpoff ( 1980a, 1989a) has identified within the Europe Neanderthal sample. The Vindija material would not therefore , be indicating a transition, but would be demonstrating the continuation of pre-existing developmental trends . The trend for thinner , less projecting, supraorbital tori with the beginnings of the separation of the superciliary arch from the supraorbital trigone identified at Vindija is also not the trend that would be required to produce the supraorbital form of the earlier Upper Palaeolithic male crania from central Europe such as Mladec 5 and 6. The population represented by the earlier Jebel lrhound material from north Africa could also be the source of these features in that they present a similar brow ridge pattern to that of the Vindija material (Smith et al. 1989) and occipital buns (cf. Chapter 3). Smith (1992) however, has suggested that these features on the Jebel Irhoud material may be the result of gene flow from Neanderthals in Europe . In general, the Vindija hominids (G3 and Gl) are best regarded as gracile Neanderthals. Although the Kolna juvenil e right maxilla displays a slight canine fossa the remainder of its morphology and that of the parietal from the site (and possibly from the same individual) display a Neanderthal morphology (Jelinek 1966, 1969, 1980c, 1982; Smith 1982, 1984). Also, as demonstrated by material from Atapuerca (Gran Dolina) and Steinheim , the presence of a canine fossa does not necessarily indicate a totally modern cranial morphology. The remainder of Wolpoffs (1980a, 1989a) transitional specimens consist of dental remains . Due to a large overlap 158

and Euclidian distance. Hahnofersand was positioned closest to La Ferrassie 1 and Amud 1. However, on a number of these analyses Hahnofersand was positioned closer to the anatomically modern Upper Palaeolithic material than to most of the Neanderthals (Brauer 1980a). Also, overall size appears to be influencing many of these analyses.

in dental metrical and morphological ranges of variation it is very difficult to ascertain if isolated teeth belonged to Neanderthals or anatomically modem humans. Grotte du Renne (Arcy-sur-Cure) has produced hominid remains from both Mousterian and Chatelperronian levels ( Oakley et al. 1971; Movius 1969). The teeth from the Chatelperronian layers display morphological continuity with those from the Mousterian layers (Mann and Trinkaus 1973; Movius 1969; Stringer et al. 1984). The juvenile temporal bone has been shown to come from a Neanderthal child (Hublin et al. 1996). There is no indication of a transition at this site and it is consistent with Saint-Cesaire in demonstrating that Neanderthal remains are associated with the Chatelperronian.

Although the frontal squama of Hahnofersand is very thick and quite flat, its overall metrical and morphological characteristics can be matched by the robust, early anatomically modern crania from central Europe such as Mladec 5, Pavlov 1 and Podbaba (Smith 1982, 1984; Stringer et al. 1984). Brauer (1980a, 1980b, 1981) did not include these individuals in his analyses. Other central European hominids such as Sala 1 have also been identified as transitional specimens (Smith 1882). Sala 1 is the frontal bone of a relatively small and probably young adult female individual with an overall Neanderthal morphological pattern, but with less rounding of the orbital margin and slight lateral thinning of the supraorbital torus (Sladek et al. 2002; Smith 1992). The dating of Sala 1 could be late Upper Pleistocene (oxygen isotope stage 3) or more probably last interglacial (oxygen isotope stage 5e )(Sladek et al. 2002), which would preclude the identification of Sala 1 as transitional unless it was transitional from Middle to the Upper Pleistocene hominids. Also, if Sala 1 is of last interglacial age, it supports the contention that lateral thinning of the supraorbital torus as identified on the Vindija material and Saint-Cesaire is the continuation of a Neanderthal morphological trend unrelated to the origin of anatomically modern humans.

The anterior teeth from Hortus and in the Sipka mandibles are said to display reduction, especially in their breadths. All of the teeth are, however, still within the Neanderthal range (Frayer 1978; Smith 1982, 1984). The Sipka incisors are outside of the Predmost range and above the Upper Palaeolithic mean (Frayer 1978; Smith 1982, 1984). The immature Hortus 4 mandible (de Lumley 1973) was used in the analysis of mandibular non-metric traits. It was well separated from both the Neanderthal and anatomically modern mandibles (Fig. 5:16). Due to its receding symphyseal region without a well-developed chin, apparent lack of retromolar gaps and the position of multiple mandibular foramina between the fourth premolar and the first molar it grouped with the Mauer and Malarnaud 1 mandibles. It is evident that none of the material that Wolpoff ( 1980a, 1981, 1989a; W olpoff et al. 1981; Frayer et al. 1993) proposes as transitional specimens can be regarded as such. Also, apart from Saint-Cesaire, which is Neanderthal in overall morphology, the remainder of the material is either very fragmentary, isolated dental material, or belonged to immature individuals.

It would be difficult to identify "transitional specimens", if they did exist due to morphological differences caused by age, sex and individual variation as well as preservation of the skeletal material and continuing problems with establishing secure dates for much of the material. However, as has been suggested by Stringer (1985; Stringer et al. 1984), there currently does not appear to be any transitional specimens in Europe.

The Hahnofersand frontal from northern Germany has also been identified as a hybrid or transitional specimen (Brauer 1980a, 1980b, 1981; Smith 1982). An amino acid racemisation date of 36,000 years BP (Fra-a-29) and a radiocarbon date of 36,000+/-600 years BP (Fra-24) have been obtained from this specimen (Brauer 1980a, 1980b, 1981 ), but there may be problems with these dates (Stringer 1989b; Stringer et al. 1984). The Hahnofersand frontal is largely complete except for the lateral and inferior portions of the brow-ridges. The frontal squama is very thick, broad and quite flat. Brauer (I 980a, 1980b, 1981) regards the contour of Hahnofersand as being similar to that of the Neanderthals, but there can be difficulties in orienting isolated frontals. The morphological features of the specimen, however, align it with anatomically modem humans. For example, the large arched brow-ridges appear to separate into medial and lateral segments and the frontal sinuses extend superiorly into the frontal squama (Brauer 1980a, 1980b, 1981; Smith 1984; Stringer et al. 1984).

Arguments for a "transition" have also focused on identifying the persistence of Neanderthal features on early anatomically modern hominids. As articulated by Frayer, assumption is that If replacement accounts for the change, the samples succeeding Middle Palaeolithic groups should not show features considered as uniquely characterising the earlier period. (1992: 180) Central Europe has provided a larger chronologically early anatomically modern sample than western Europe. The western European material is also generally more gracile than the central European hominids. It has been argued that that the central European early anatomically modern human sample (such as the individuals from Mladec, Predmost, Brno and Stetten) displays Neanderthal reminiscent features that can only be accounted for by advocating an in situ transition (Jelinek 1969; Frayer 1992; Frayer et al. 1993; Smith 1982, 1984, 1985, 1992; Smith and Paquette 1989; Smith and Ranyard 1980; Smith et al. 1989; Wolpoff 1980a, 1989a; Wolpoff et al. 2001 ).

Brauer ( 1980a, 1980b, 1981) carried out a number of multivariate analyses utilising the Hahnofersand frontal and Neanderthal and anatomically modem crania. He used principal components analysis, principal co-ordinates analysis, cluster analysis, Penrose size and shape distances 159

There are few late central European Neanderthal postcranial remains , but the early anatomically modem human postcranial sample from the region does not regularly reveal Neanderthal reminiscent features (e.g. Predmost 14 has a dorsal axillary scapular border form; Jelinek 1969; Smith 1982, 1984; Stringer et al. 1984). Frayer (1992) has shown that the majority of Neanderthals have the dorsal axillary scapular border form, whereas early European Upper Palaeolithic material generally has the bisulcate axillary scapular border form. There is also no evidence of the muscular hypertrophy of the Neanderthals and the distal limb segments are longer , suggesting an adaptation to a warm climate and/or a modern locomotor pattern (Trinkaus 1981, 1983, 1986, 1989).

supra-1ruac fo ssa on Neanderthals ( 100%), early Upper Palaeolithi c (25%) and late Upper Palaeolithic (23.7%) hominids , but acknowledges that the form of the suprainiac fossa on the anatomically modem material is different to that seen on the Neanderthals . Also a suprainiac fossa can be found on earlier material from Africa (Eliye Springs 11693 and possibly Kabwe 1) and western Asia (Skhiil 5), but again these are of a different form to that of the Neanderthals. A unique combination of morphological features characterise the Neanderthals , not isolated individual features, especially when these features are not found on all Neanderthals or can be found on earlier hominid material from other geographical regions . W olpoff et al. (2001) used pairwise difference analysis to analyse non-metric traits so as to compare the male Mladec 5 and 6 individuals to Neanderthal crania and the Skhiil and Jebel Qafzeh hominids (Skhiil 5, 4 & 9 & Qafzeh 6 & 9). Based on the 30 non-metric traits used , Mladec 5 showed less pairwise differences to Skhfil 4, Skhiil 9 and Jebel Qafzeh 6 than to the Neanderthal crania examined and Skhfil 5 and Jebel Qafzeh 9 (Wolpoff et al. 2001 :Table 2) . Based on the 22 non-metric traits used, Mladec 6 showed less pa irwise difference s to the Neanderthal crania examined than to the Skhiil and Jebel Qafzeh material (Wolpoff et al. 2001 :Table 3). Wolpoff et al. (2001 ) indicated that Mladec 5 had a Neanderthal-like sagittal contour , a well-developed occipitomastoid crest, limited mastoid projection and evidence of midfacial prognathism , while Mladec 6 had a Neanderthal-like elliptical suprainiac fossa, lambdoidal flattening and a short posterior face on the occiput. Wolpoff et al. (2001 :296) concluded that "a dual-ancestry model cannot be rejected" for Europe and that there is no evidence to suggest that ... the Mladec males can be grouped with the Levantines to the exclusion of European Neandertals. (2001 :296) It is worth noting that the two robust Mladec males had varying patterns of differences/similarities in these analyses with Mladec 5 less similar to the Neanderthals and more similar to the Skhiil and J ebel Qafzeh hominids than Mladec 6.

The central European early anatomically modern sample , especially the males , is very robust. They have large brow ridges of modem type in that they can be separated into medial and lateral segments and so do not form a torus in the Neanderthal sense. Occipital buns are commonly found on the early modern sample from central Europe, but these are higher on the cranium than the Neanderthal "chignon" and do not position the nuchal plane as horizontally (Smith 1982, 1984). Smith (1984 ) refers to these structures as "hemibuns" . Also , the presence of an occipital bun cannot be regarded as a Neanderthal reminiscent feature for central European anatomically modem humans because the preceding central European Neanderthals do not appear to have had occipital "chignons" (Smith 1976b, 1982, 1984; Wolpoff et al. 1981). This structure is more typical of western European Neanderthals. However , it is not common on early anatomically modem humans from western Europe. Cro-Magnon 3 does have a pronounced occipital bun, which Wolpoff (1980a) identified as ''Neanderthal-like" , but the morphology is different to that of the Neanderthals (Gambier 1989). This feature on Cro-Magnon 3 may also be related to an unusually late, posteriorly-directed growth of the brain (Trinkaus and LeMay 1982), as demonstrated by the occurrence of numerous extra sutural bones in the lambdoidal suture , and not due to genetic input from the Neanderthals. Some studies have focused on the frequency of Neanderthal morphological features on early anatomically modem hominids (for example Frayer 1992; Smith 1978). Based on this type of data, Frayer et al. argued the Evidence represented in the fossil record of late Pleistocene Europe demonstrates a persistence of so-called unique Neanderthal features over time, bridging the populations of the Mousterian and Upper Paleolithic . (1993:28)

However , pairwise difference analysis is generally used for analysing genetic sequence data and so its appropriateness for analysing continuous non-metric traits on hominids as done by W olpoff et al. (2001) has not been established . The choice of non-metric traits and hominids included could also influence the outcomes of the analyses. Had the two female crania from Mladec been included in the pairwise difference analyses the results may have been different as these more gracile individuals have few Neanderthal-like morphological features and provide little evidence for continuity. Finally, Wolpoff et al. (2001) contend that "visual inspection" readily supports closer links between Neanderthals and early European Upper Pala eolithic material. In support of this contention Wolpoff et al. (2001) provided illustrations of Spy 2, Mladec 5 and Jebel Qafzeh 9 in lateral views. It is worth noting that elsewhere Wolpoff (W olpoff and Caspari 1990) has argued that Jebel Qafzeh 9 should not be used to typify the Jebel Qafzeh hominids as it is an adolescent and appears more modem than Jebel Qafzeh 6 in that it has a higher and more rounded cranial vault, lacks lambdoidal flattening and has a reduced supraorbital region. Jebel Qafzeh 6 was the

However, the individual morphological features are either not unique to the Neanderthals or not found on all Neanderthals . For example , both Smith (1978) and Frayer ( 1992) use the presence of the horizontal-oval mandibular form of mandibular foramen , which they have identified on 46.2% (Smith 1978) or 52 .6% (Frayer 1992) of Neanderthals and 23% of Upper Palaeolithic anatomically modem hominids (Smith 1978), or 44.4% of early Upper Palaeolithic and 5.3% of late Upper Palaeolithic material to argue for continuity from Neanderthals to early anatomically modem Homo sapiens, yet only around half of the Neanderthals have the feature. Frayer ( 1992) also details the presence of a 160

which display a unique and quite specialised morphological pattern (Table 5:6; Howells 1974; Hublin 1984; Rak 1986; Stringer 1985, 1989a, 1989b; Stringer et al. 1984).

most similar individual to both Mladec 5 and 6 on these analyses (Wolpoff et al. 2001:Tables 2 & 3). An alternative interpretation would have occurred from "visual inspection" if one of the female Mladec crania had been included in the illustration.

It must also be remembered that there are robust anatomically modem hominids in western Asia and Africa between 50-100,000 years ago, when Neanderthals were still developing their unique morphology in Europe.

Even if one were to accept the contention that the presence of these Neanderthal-like features on Upper Palaeolithic hominids reflected gene flow between Neanderthals and robust anatomically modem Homo sapiens, this would only indicate some degree hybridisation between the two groups, not that Neanderthals evolved into or had any significant input into the origin of anatomically modem Homo sapiens.

Conclusion As has been outlined, there are two major explanations for the origin of anatomically modem humans in Europe: t~e Multiregional Hypothesis, and the Replacement Hypothesis. The former hypothesis attempts to identify transitional specimens and to document the transition from the Neanderthal to the anatomically modem morphological pattern. The latter hypothesis highlights the morphological dichotomy evident in Europe and proposes that the anatomically modem morphological pattern has its origin outside of Europe.

As previously discussed, there do not appear to be any "transitional" specimens in Europe that document a morphological change from Neanderthals to anatomically modem humans. What we have in central Europe is gracile Neanderthals, such as the Vindija material, and very robust anatomically modem crania, especially the males. Stringer et al. ( 1984) argued that the robust anatomically modem humans from central Europe resemble the Jebel Irhoud hominids from north Africa more than they resemble the Neanderthals in their archaic features.

There are some morphological changes in the Neanderthal remains, especially those from central Europe . However, the documentation of a trend towards gracilisation within the Neanderthals does not prove that a transition did occur.

No anatomically modem hominid cranium from Europe displays a predominance ( or even a large number) of typical Neanderthal features. The anatomically modem early Upper Palaeolithic crania are robust and in general share only primitive features, such as large brow-ridges, prognathic faces and moderately low vaults, with the Neanderthals. Frayer, who supports regional continuity in Europe, has stated in regards to the male crania from Mladec

An ordered similarity matrix analysis was undertaken to ascertain if there had been an orderly morphological progression in Europe, or if there was evidence of a major discontinuity within the sequence (Tables 5:7 and 5:8). The hominid sample used, which spans the mid-Middle Pleistocene through to the late Upper Pleistocene , comes predominantly from western Europe. The Arago reconstruction, which utilises Arago 21, Arago 4 7 and the Swanscombe occipital was also included in the analysis ( 1er Congr. Intern. Paleont. Hum., Nice 1982, Pretirage, Vol. 1). The gradual evolutionary development of the European Neanderthals is evident from this analysis (Table 5:8). The Middle Pleistocene hominids were shown to display strong similarities to the Upper Pleistocene western European Neanderthals . This group is clearly differentiated from the western European anatomically modem humans used in the analysis (Table 5:8). The ordered similarity matrix analysis has identified a major disconformity in the hominid sequence, and so supports the Replacement Hypothesis, for western Europe at least. Similar results were obtained by Waddle ( 1994) using matrix correlation methods to compare morphological distances and Harvati (2003) who found there were greater intra-specific differences between Neanderthals and Late Pleistocene hominids and modem humans than was evident between recent human populations.

...each exhibit a number of archaic features which link them to earlier Neanderthal populations. (1986:251) The pattern however is not consistent. Mladec 5 for example, displays a left mastoid region that recalls the Neanderthals, whereas on the right side it resembles anatomically modem crania (Frayer 1986). There is also considerable morphological variation between male and female crania from the early Upper Palaeolithic. As Frayer observed ...the differences between Neanderthal females and the two female crania from Mladec are striking and provide little evidence for continuity between the two samples. (1986:249) The point that needs to be remembered is that a large browridge, low vault, prognathic face and protruding occipital region does not make a Neanderthal. The characteristic Neanderthal features of the splanchnocranium and/or occipitomastoid region (Table 5:6) are not found in combination on anatomically modem humans from Europe. This sample displays an "en maison" transverse contour in norma occipitalis, brow-ridges with definable medial and lateral segments, moderate to slight mid-facial prognathism, generally large projecting mastoid processes without occipitomastoid crests (although Wolpoff [1989a] contends that some of the early anatomically modem crania from central Europe have non-projecting mastoid processes) and moderately developed mental eminences. This morphological pattern is clearly distinct from that of the Neanderthals,

The Earliest Anatomically Modern Humans in Europe Refinements in the dating of sites throughout Europe have focused attention on the possible overlap of Neanderthals and anatomically modem humans. If anatomically modem humans replaced the Neanderthals one would expect some chronological overlap between the two groups. In western Europe late Neanderthals have been dated at 36,300 years BP at Saint-Cesaire (Mercier et al. 1991), 30,000 years BP at 161

Table 5:7. Measurements used in the Ordered Similarity Matrix for Europe.

Glabella-occipital length Nasion-occipital length Glabella-inion length Basion-nasion length Basion-bregma height Auricula-bregma height Maximum cranial breadth Maximum and minimum frontal breadth Bistaphanic breadth Biauricular breadth Biasterionic breadth Basion-prosthion length Nasion-prosthion height Nasal height and breadth Orbit height and breadth (left) Palate length and breadth Palate corpus thickness between left Ml and M2 Premolar chord between left PM3 and M2 Prosthion -end of dental arch Biorbital breadth (ectoconchion) lnterorbital breadth ( dacryon ) Simotic chord Cheek height (left) Nasion-bregma (frontal) chord and arc Bregma-lambda (parietal) chord and arc Lambda-opisthion (occipital) chord and arc Lambda-inion chord and arc Nasion-opisthion arc Transverse arc via bregma

Table 5:8. Ordered Similarity Matrix for Europe. For details on the crania used see Table 5:2. Q=328, Q MAX = 1320, Q/Q MAX=.248485 Crania p A21&47 St Sal Gl LCI LFl

cc

CMI Ch LP5

p

--

A21&47 # * $ $ $ $ $ $ + +

--

--

* # $ $ $ $ $ +

--

Sal

St

Gl

$ $ * $ $ $ $ + +

$ $ $ * # $ #

$ $ $ # * $ $

+

+ +

---

-+

+

--

+

LCl $ $ $ $ $ * $ +

----

cc

LFl

CMl

Ch

+

--

--

$

+ +

+

$ $ $ # $ $ *

+ + + + +

+ + +

* $ $

+

--

+

+

--

LP5 --

---

--

+

--

+

+

+

--

--

+

+ $ + * $

---

$ *

+ + $ *

Key: *=Identical, #=Very Similar , $=Similar, +=Not Very Similar, --=Unlike Note: See Table 5:2 for names of crania abbreviated above.

Zafarraya (Hublin et al. 1995; Mellars 1998a; Sanchez 1999) and at Arcy-sur-Cure to 33,800 years BP (Hublin et al. 1996; Smith et al. 1999) and in Central Europe to 28,000 years BP at Vindija (Smith et al. 1999) . These dates demonstrate that Neanderthals were present in Europe while anatomically modem humans were present in western Asia and Africa (cf . Chapters 2, 3 and 4), and infer that the Neanderthals were contemporary with anatomically modem humans in Europe.

Establishing early dates for the remains of anatomically modem humans in Europe has however, proven difficult. Smith et al. ( 1999) provide a good discussion of the claims of the earliest dated anatomically modem human remains within Europe and conclude that many of the early dates and the stratigraphic provenances of the hominids can be questioned or the remains are not diagnostic. The V elika Pecina frontal from northwestern Croatia , had been regarded as the earliest conclusively dated anatomically modem hominid in Europe because a date of 162

As discussed in Chapter 4, anatomically modem humans are associated with a Levalloiso-Mousterian industry at Skhul and Jebel Qafzeh in western Asia.

33,850+/-520 years BP (GrN-4979) had been obtained from an Aurignacian level above the layer from which the hominid frontal bone was said to have been recovered (Smith 1976a, 1982, 1984). However, a direct accelerator mass spectrometry (AMS) radiocarbon date of 5,045+/-40 years BP (OxA-8294) has been obtained from the Velika Pecina frontal (Smith et al. 1999). The hominid frontal must therefore be intrusive.

Cranial fragments and some postcranial material that is morphologically aligned with early anatomically modem Homo sapiens have been recovered from the island of Crete (Facchini and Giusberti 1992). A calcareous breccia in direct contact with the hominid bones has been chronometrically date by the protactinium/uranium method to 51,000+/-12,000 years BP (Facchini and Giusberti 1992). If the date of the hominid material can be confirmed it would make the Crete hominid the earliest anatomically modem specimen found in Europe.

Jelinek (1969) placed the Neanderthal mandible from Sipka and the anatomically modem hominids from Mladec within the Podhradem interstadial and regarded them as being contemporary with each other. As alluded to earlier, there are major problems with the establishment of this interstadial, in that it may include two separate interstadials corresponding to the western European Hengelo and Denekamp/Arcy interstadials (Smith 1982; Stringer 1982; Valoch 1968; Woillard and Mook 1982). Smith (1984) placed the Sipka mandible within the "Lower Wiirm" stadial (following Valoch 1968), which he dates between 38,000 and 45,000 years BP. The Mladec hominid material probably belongs to the "Wurm II/III" interstadial and so would be younger than 40,000 years BP. The Mousterian associated Sipka mandible and the probably Aurignacian associated hominids from Mladec do not appear, on current evidence, to have been contemporary .

The Mladec hominids may be older than 30,000 years BP (Frayer 1986; Jelinek 1969; Smith 1982, 1984; Wolpoff 1980a). The Predmost hominid sample is most probably contemporaneous with the Pavlovian/Eastern Gravettian occupation at the site (Smith 1984), which has provided a date of 26,320+/-240 years BP (GrN-6852). Other central European anatomically modem hominids that date to at least 25,000 years BP come from Paderbom, Brno, Dolni Vestonice and Pavlov (Henke and Xirotiris 1982; Jelinek 1969; Smith 1982, 1984). Germany has produced a number of relatively early anatomically modem hominids. As previously discussed, the Hahnofersand frontal has provided an amino acid racemisation date of 36,000 years BP (Fra-a-29) and a radiocarbon date of 36,300+/-600 years BP (Fra-24), but there appear to be problems with these dates (Brauer 1980a, 1980b, 1981; Stringer 1989a, 1989b; Stringer et al. 1984). Dates of approximately 30,000 years BP have been obtained for Aurignacian levels at the site of Stetten from which an anatomically modem human calvaria has been recovered (G. Brauer pers. comm.; F.H. Smith pers. comm.). A calvaria from Kelsterbach has a radiocarbon date of 31,200+/-600 years BP (Fra-5) and an amino acid date of 32,000 years BP (Henke and Xirotiris 1982; Smith 1984). The relatively gracile nature of this find, when compared to the other early Upper Palaeolithic material from the region, has resulted in its date being questioned (Smith 1984; Wolpoff 1989a). The geological position of the find, however, suggests that it is at least 21,000 years old (Smith 1984).

The anatomically modem Vetemica calvaria was initially attributed to a Mousterian level at the site (Coon 1962). A date of greater than 43,200 years BP (GrN-4984) has been obtained from the layer immediately below the level from which the hominid was thought to be derived (Smith 1976a, 1977, 1982; Vogel and Waterbolk 1972). Reinterpretation of the stratigraphical context of the hominid indicates that it was an intrusive burial and so is probably of Aurignacian age, at the most (Smith 1976a, 1977, 1982; Stringer and Burleigh 1981). Smith (1976b) suggested that the morphology and lack of fossilisation of the calvaria align it with the later, mostly Mesolithic, skeletal remains from the site. There are no indubitable Mousterian-associated anatomically modem humans in central or western Europe. At the site of Carigiiela (Pillar) near Granada in Spain, anatomically modem remains (a parietal fragment, mandible and tibia) have been recovered from layers that are said to contain what is referred to as a MousterioAurignacian industry or a Mousterian industry with Aurignacian influences (Garcia 1960; de Lumley and Garcia 1971; Oakley et al. 1971; Stringer et al. 1984). Fluorine and nitrogen analyses, although generally inconclusive, allow the possibility that this material is not contemporary with the animal bone tested (Oakley 1980; Oakley et al. 1971; Oakley [ 1980] indicates that all three hominids were recovered from level 2, whereas Garcia [1960] stated that one of the remains came from level 3, which is the same level as the animal bone tested). There is the possibility that the hominids were burials into the Mousterian levels. It is also possible that the levels from which the hominids were recovered were mixed and so the hominid remains may not be Mousterian associated even if they belong to these levels.

Two Austrian sites have also produced the remains of early anatomically modem hominids. Miesslingtal has provided an Aurignacian associated, fragmentary, immature mandibular corpus, while W illendorf has provided an Eastern Gravettian associated symphyseal segment of a mandible (Oakley et al. 1971; Smith 1984). The earliest anatomically modem hominids from western Europe, such as the Cro-Magnon remains, appear to be younger than 30,000 years BP, since they are generally associated with evolved Aurignacian industries (Oakley et al. 1971; Stringer et al. 1984; Wolpoff 1980a). The Combe Capelle skeleton was recovered from a Chatelperronian (Perigordian 1) layer, although it now appears as though the skeleton was an intrusive burial from higher Aurignacian levels (ApSimon 1980; Stringer et al. 1984) or even post-Palaeolithic levels (Gambier 1989). 163

There are a number of fragmentary hominid remains that could represent either late Neanderthals or early anatomically modern humans .

Vogelherd Kelsterbach Kent's Cavern

The Vindija Gl and F complex hominid remains have already been discussed. The F complex remains are associated with an Aurignacian industry, whereas there are major problems with the attribution of the G 1 artefacts to the Aurignacian .

31,900+/-l , 100 years BP 31,200+/-600 years BP 32,000 years BP 30,000 +/-900 years BP

The preceding discussion and dates indicate that sites with Neanderthal remains and sites with the remains of anatomically modern humans can be dated to the same chronological period , but do not conclusively demonstrate that Neanderthals and moderns were contemporary in the same areas/regions within Europe .

Dental remains have been recovered from the Grotte du Renne (Arcy-sur-Cure) associated with Mousterian , Chatelperronian (a temporal bone was also recovered) and Aurignacian industries (Oakley et al. 1971). Dates of 33,860+/-50 years BP (GrN- 1742) and 33,500+/-400 years BP (GrN-1736) have been obtained from Bed 8, one of the Chatelperronian layers (Movius 1969; Oakley et al. 1971; Vogel and Waterbolk 1963). An accelerator mass spectrometry radiocarbon date of 33,820+/-720 years BP has also been reported for the Chatelperronian at the site (Smith et al. 1999). A date of 30,800+/-250 years BP (GrN- 1717) has been obtained from Bed 7, an Aurignacian layer (Oakley et al. 198 1).

Bocquet-Appel and Demars (2000) undertook an analysis of the radiocarbon dates from sites around the "transition" and mapped the "contraction" of the Neanderthals and the "expansion " of modem humans. They found that before 40,000 years BP Neanderthals were present throughout Europe, but over the next 5,000 years "absence zones" develop (Bocquet-Appel and Demars 2000 :Figure 7). From 35-30,000 years BP Neanderthal presenc e is restricted to southern France and the western third of the Iberian peninsula and they disappea r by 27,500 years BP (Appel and Demars 2000:Figure 7). Anatomically modern humans were found to be absent from Europe at 40,000 years BP (Appel and Demars 2000:Figure 8). Over the next 5,000 years anatomically modern humans are present throughout eastern Europe and continue to spread westward during the period to 30,000 years BP, when there is only a small area of absence in the western Iberian peninsula (Appel and Demars 2000:Figure 8). By 27,000 years BP anatomically modem humans have colonised all of Europe (Appel and Demars 2000:Figure 8). Pettit and Pike (2001) question the statistical methods used by Appel and Demars study and outline problems with the accuracy of radiocarbon dating during this period . They concluded ... chronological reconstructions of the palaeodemography of Neanderthal extinction and modem human expansion are premature at best, and possibly seriously misleading . (2001 :417)

Two isolated deciduous molars have been recovered from an Uluzzian level at Cavallo Cave in Italy, and are dated in excess of31 ,000 years BP (Stringer et al. 1984). The Bulgarian site of Bacho Kiro has produced fragmentary hominid remains from levels containing an Aurignacian-like industry (Kozlowski 1982). The hominid material consists of a fragment of the right corpus of a mandible and a right upper canine from Layer 6a/7, a right mandibular central incisor from Layer 7/6b, a fragment of the right parietal and a right lateral mandibular incisor from Layer 7, and a fragment of the left mandibular body containing a single worn deciduous first molar from Layer 11 (Glen and Kaczanowski 1982). Radiocarbon dates have been obtained for these levels (Kozlowski 1982). These dates are 29,150+/-950 years BP (Ly-1102) for Layer 6a/7, 32,700+/-300 years BP (GrN-7569) for Layer 6b, and greater than 43,000 years BP (GrN-7545) for Layer 11 (Kozlowski 1982). Glen and Kaczanowski (1982) suggested that most of the permanent teeth from Bacho Kiro were similar to N eandertal teeth. Ginter and Kozlowski (1982) observed that some features of the dentition suggested a transitional position between the Neanderthals and Homo sapiens , although they favoured a classification of the Bacho Kiro material as Homo sapiens . The Bacho Kiro hominids are often referred to as primitive modern Homo sapiens (Howell 1984; Smith 1982, 1984; Stringer et al. 1981; Stringer et al. 1984). The material , however , is too fragmentary for a clear identification of it as either Neanderthal or anatomically modern.

Litbic Material A discussion of the lithic traditions within Europe can assist in determining if there was continuity or if there was replacement in Europe . Mousterian industries are found throughout Europe from at least the last interglacial through the first half of the last glacial down to "W firm III" (Table 5: 1; Bordes 1968; Dennell 1983a, 1983b; Mellars 1986, 1989a, 1989b; Mellars et al. 1987), that is, from oxygen isotope stage 5 to oxygen isotope stage 3 (Table 5: 1; Dennell 1983a, 1983b; Griin et al. 1997).

Recovered from an Aurignacian layer dated to 30,900+/600 (GrN-1935) at the Hungarian site oflstallosko was the crown of an unerupted mandibular second molar (Fig. 5:5; Howell 1984; Oakley et al. 1971; Smith 1982, 1984).

Based on the percentages of tool types Bordes ( 1961, 1971, 973; Bordes and Sonneville-Bordes 1970) identified four major divisions within the Mousterian technocomplex of southwestern France: Typical, Denticulate , Charentian (including Ferrassie and Quina variants) and Mousterian of Acheulian Tradition (including Types A and B). Bordes ( 1961, 1971, 1973; Bordes and Sonneville-Bordes 1970) did not identify major temporal changes in these divisions, which Bordes thought were being produced at the same

Smith et al. ( 1999) provided the following dates for other early anatomically modern human remains from Europe:

164

time by distinct ethnic groups. Sedimentological studies of the caves of southwestern France and Cantabrian Spain have supported Bordes' conclusion that the various divisions were contemporary (Butzer 1981; Laville 1973; Laville et al. 1980). This patterning, however, is not found outside this region (Bordes 1968; Dennell 1983a). It was suggested that the Mousterian variants were manufactured by contemporaneous, but different, cultural groups (Bordes and Sonneville-Bordes 1970). However, it is difficult to explain how and why five "cultural groups" could have coexisted in southwestern France and northern Spain for such a long period of time (Dennell 1983a; Shackley 1980).

Waterbolk 1972). This industry is generally regarded as being the product of an acculturation process at the junction of the Middle and Upper Palaeolithic {AllsworthJones 1986a, 1986b; Kozlowski 1990; [definition of acculturation as per Mellars 1989b:353]). The Szeletian is also found in the same region in which early Aurignacian industries have been found. No hominid remains are conclusively associated with the Szeletian. At Dzerava Skala (Palffy) in eastern Slovakia the germ of a lower left second molar was recovered in a layer that contained Szeletian and Aurignacian components mixed by cryoturbation, and at Silicka Brezova a lower molar, although not in direct association with any lithic material, may have been associated with the Szeletian (Smith 1982, 1984). The best candidates may be the fragmentary hominid material from level G 1 at Vindija. As noted earlier, the limited lithic material from G 1 includes pieces typical of both the Mousterian and the Upper Palaeolithic associated with three massive-base bone points and a splitbased bone point (Karavanic and Smith 1998). The level G 1 material culture has been regarded as both Aurignacian and Szeletian (Karavanic and Smith 1998; Malez et al. 1980; Smith 1982, 1984; Wolpoff et al. 1981).

Mellars (1969, 1973) has argued that there was change through time within the Mousterian and he placed Bordes' various divisions in a temporal sequence from Typical/Denticulate through Ferrassie, Quina and Denticulate/Typical to Mousterian of Acheulian Tradition. Mellars (1986, 1989a) has used thermoluminescence dating at Le Moustier (Valladas et al. 1986) to support his arguments.

It has also been suggested that the divisions identified by Bordes could have been caused by adaptations to changing environments and/or different tool kits being used for seasonally or functionally different activities (Binford 1973; Binford and Binford 1966, 1968).

The Chatelperronian, which probably dates from between 38,000 and 33-31,000 years BP {ApSimon 1980; Harrold 1983, 1989; Mellars 1998a, 1998b, 1999 [especially 1999:Fig. 1, Table 1]; but see d'Errico and Sanchez Gofu 2003; d'Errico et al. 1998; Zilhao and d'Errico 1999, 2000 for an earlier > 35-40,000 years BP dating scheme), occupies a similar position in France to that held by the Szeletian in central Europe. Bordes stated that the ...lower Perigordian [Chatelperronian] is derived from a local development of an Acheulian-tradition Mousterian in the Upper Palaeolithic. (1968: 148)

Mousterian variation in southwestern France and northern Spain remains a very complex matter and all of the explanations outlined have problems. Future attempts at explaining this variation must take into consideration temporal and functional variation as well as raw material differences and reduction strategies (Dibble 1983; Rolland 1981). Many discussions of the industries involved in the transition from the Middle to Upper Palaeolithic in Europe are available (cf. Ailsworth-Jones 1986a, 1986b; Ashton 1983; Bricker 1976; Bordes 1968; Coles and Higgs 1969; Collins 1986; Dibble 1983; Frayer 1978; Kozlowski 1990; Harrold 1983, 1989; Mellars 1973, 1989a, 1989b, 1998b; Pradel 1966; Straus 1983; Svoboda 1984, 1986, 1989; Valoch 1968, 1986; White 1982, 1983; Wymer 1982; and references therein) and so a detailed discussion of the nature of each industry will not be undertaken here.

Harrold (1983, 1989) contends that the Chatelperronian did not develop gradually out of the Mousterian, but was an abrupt development from the Mousterian (Mousterian of Acheulian tradition) under the influence of the Aurignacian. He regarded it as a heavily acculturated derivation of the Mousterian, (see also Mellars 1989b, 1998a, 1998b, 1999). Harrold ( 1983, 1989) also identified time-transgressive replacement of the Chatelperronian by the Aurignacian from the south and east towards the north and west (see also Lynch 1966). Zilhao and d'Errico, based on an early dating scenario (>35-40,000 years BP), have argued that Chatelperronian industries and the associated bone tools and ornaments represent an independent autogenous development and not acculturation, but this remains a minority view (d 'Errico and Sanchez Gofii 2003; d'Errico et al. 1998; Zilhao and d'Errico 1999, 2000; see also discussion in Bahn 1998, Mellars 1998a and Wong 2000).

Laville (1973; Laville et al. 1980) has suggested that the Middle to Upper Palaeolithic transition in southwestern France was marked by an erosional episode. This could explain the lack of clearly transitional Middle/Upper Palaeolithic industries in western Europe. There is a group of industries found throughout Europe, including the Szeletian, Chatelperronian and Uluzzian that combine flake and blade technologies. These industries have limited distributions (Mellars 1989b:Fig. 3) and chronologically overlap and are interstratified with layers containing early Aurignacian industries at some sites.

The Chatelperronian levels at Grotte du Renne (Arcy-surCure) seem to demonstrate that Neanderthals did make ornaments (eg. pierced tooth pendants and ivory rings) and bone tools (d'Errico et al. 1998; Zilhao and d'Errico 2000). This may indicate that contact between the Neanderthals and anatomically modem humans initiated the start in the use (Neanderthals) or the increased use (anatomically modem humans) of symbols of group

The Szeletian dates from between possibly 43,000 and 33,000 years BP and appears to have developed out of the local Mousterian in central Europe (Ailsworth-Jones 1986a, 1986b; Smith 1982; Valoch 1968, 1986; Vogel and 165

identity. However, artefacts or objects (eg. national flags) that are being used to identify a group or mark cultural, social or territorial boundaries are usually markedly different in style (Franklin 1986, 1989). Due to the similarity in general appearance to Aurignacian ornaments ( as opposed to differences in manufacturing methods) the ornaments may not be being used by these particular Neanderthals as a conscious symbol of group identity, but rather may be reflecting a degree of social interaction between the two groups (Franklin 1986, 1989).

sites such as La Rochette , Roe de Combe, Abri Pataud, Grotte du Renne (Arcy-sur-Cure), Les Cottes E, La Quina, Le Flageolet I and La Ferrassie (d'Errico and Sanchez Gofii 2003; Frayer 1978; Mellars 1998b, 1999; Mellars et al. 1987; Movius 1972, 1977; Oakley et al. 1971; Vogel and Waterbolk 1963, 1967, 1972). The Aurignacian in Germany and Austria appears to be earlier than in France with dates of 45-35,000 years BP obtained from sites such as Willendorf, Geissenklosterle Hohlenstein-Stadel , Vogelherd and Krems-Hundssteig (Allsworth-Jones 1986b; Conard and Bolus 2003; Howell 1984; Mellars 1998b; Valoch 1986; Vogel and Zagwijn 1967). As in France , the Aurignacian does not appear to have clear antecedents in Germany or Austria (Bordes 1968; Frayer 1978; Wymer 1982).

Dating of Mousterian and Aurignacian occupation in the Iberian Peninsula appears to demonstrate a considerable period of coexistence in neighbouring regions (Mellars 1989b, 1998a, 1998b, 1999). In Portugal and southern Spain sites with Mousterian industries have been dated to 30-35 ,000 years BP, whereas in Spain north of the Ebro River Aurignacian industries are dated to 38-40,000 years BP (Mellars 1989b, 1998a, 1998b, 1999; but see d'Errico and Sanchez Gofii 2003 ; d'Errico et al. 1998; Stringer and Davies 2001; Zilhao and d'Errico 1999). It has been suggested by d'Errico and Sanchez Gofii (2003) that during the cold Heinrich 4 event of oxygen isotope stage 3 (35,300-33 ,900 years BP) the southern Iberian peninsular was covered by desert-steppe-like environments that not only caused a contraction of Neanderthal populations, but also may have assisted in Neanderthal persistence in this region by limiting the movement of modem humans into the region.

The earliest European Aurignacian industries occur in central Europe. There is a date of greater than 43,000 years BP (GrN-7545) for an Aurignacian-like industry in Layer 11 at Bacho Kiro (Kozlowski 1982). This layer lacks splitbased bone points, but these are found in overlying levels (Howell 1984; Kozlowski 1982; Valoch 1986). A similar industry is found at the Hungarian site of Istallosko, where the lower levels contain split-based bone points (Howell 1984; Valoch 1986). The lower levels at the site have provided dates of 44,300+/-1,000 years BP (GrN-4659) and 39,700+/-900 years BP (GrN-4658), and the upper levels have been dated to 30,900+/-600 (GrN-1935) and 31,540+/-600 (GrN-1501) , but the accuracy of some of these dates has been questioned (Frayer 1978; Howell 1984; Smith 1982; Valoch 1986; Vogel and Waterbolk 1963). Other dates of greater than 35,000 years BP for early Aurignacian industries in central Europe come from Temnata, Kovolero I, Vedrovice II, Kuparovice I, Pesko and Samuilica (Mellars 1998b; Smith 1982; Valoch 1986; Vogel and Waterbolk 1972; Vogel and Zagwijn 1967). It is worth noting that the east European plain has provided dates in the range of 39-36,000 years BP for the earliest Upper Palaeolithic (Dolukhanov et al. 2002). Even earlier dates are recorded for sites in south-central Siberia such as Kara-Tenesh where a date of 43,200+/-l ,500 years BP has been obtained for the earliest Upper Palaeolithic levels (Dolukhanov et al. 2002). Also, Mezmaiskaya Cave in the northern Caucasus has provided a date of 32,010+/-250 years BP (Beta-113536) for what is described as an early Upper Palaeolithic assemblage (Golovanova et al. 1999).

Mellars (1998a) contends that even with this period of coexistence in the southern Iberian peninsular, there is no conclusive archaeological evidence that these late Iberian Neanderthals adopted any behavioural or technological innovations from the anatomically modem humans producing the Aurignacian industries in northern Spain, as is evident elsewhere in Europe (ie. pierced tooth pendants , ivory rings and bone tools or industries similar to the Chatelperronian, Szeletian and Uluzzian). The only clearly identifiable hominid remains associated with the Chatelperronian are the Neanderthal hominids from Saint-Cesaire (ApSimon 1980; Leveque and Vandermeersch 1980, 1981) and Grotte du Renne (Arcysur-Cure)(Hublin et al. 1996), although isolated teeth have also been recovered from Chatelperronian levels at other sites (Stringer et al. 1984). As has been mentioned, the Aurignacian is often found interstratified with the Szeletian or the Chatelperronian, and it eventually replaces both of these industries (Allsworth-Jones 1986a, 1986b; Frayer 1978; Harrold 1983, 1989; Howell 1984; Svoboda 1989; Valoch 1968, 1986).

On chronological grounds, one can argue that the western European Aurignacian is derived from central Europe. However, the early dates from this region only demonstrate the occurrence of the Aurignacian. They do not document its origin. Valoch ( 1968) advocated for a local central European origin for the Aurignacian , whereas Allsworth-Jones (1986a, 1986b) and Kozlowski (1982, 1990) favour an external origin for it. However, Svoboda ( 1989) stated that the Bohunician industry , which employs levalloisian technology, appeared suddenly at around 44,000 years BP, without links to the local Moravian Mousterian , and so must represent a migration into the region. He also argues that it demonstrate s progr essive changes from the Middle to the Upper Palaeolithic. The makers of the Bohunician are unknown and could be Neanderthals or anatomically modem humans.

The Aurignacian has no antecedents in western Europe and is generally thought to have developed elsewhere. As Bordes stated ...the Aurignacian - unlike the Perigordian would not seem to have originated in western Europe, but to have arrived there ready-made ... (1968:155) In southwestern France the early Aurignacian generally dates between 33,000 and 36,000 years BP , for example, at 166

It is difficult to identify where the central European Aurignacian was originally derived. The Upper Palaeolithic in western Asia, north Africa and sub-Saharan Africa does not appear until around 40-45,000 years BP (Bar Yosef 1989; Bergman 1989; Clark 1982; Deacon 1984; Debenath et al. 1986; Ferring 1975; Smith, P.E.L., 1982), which is the same age as, or younger than, the dates for the early Aurignacian in central Europe. Howell (1984), however, contends that the early Aurignacian industry at Bacho Kiro is reminiscent of the preAurignacian of western Asia as seen at the site of Yabrud. Western Asia may therefore, have been the ultimate source of the European Aurignacian.

recent human populations. These results again demonstrate the morphological differences between Neanderthals and anatomically modem humans, which was reinforced by the results of the multivariate statistical analyses (Harvati 2003: Figs. 3, 5, 6, 8 and 9). Harvati (2003) concluded that her studies supported the view that the Neanderthals represented a different species that did not contribute to the evolution of modem humans in Europe. Overall, it appears as though the Neanderthals developed gradually in Europe, over perhaps 250,000 years, and were replaced by anatomically modem humans following a period of coexistence of possibly 10,000 years. Therefore, there is no evidence of a catastrophic end of the Neanderthals, in so far as the interstratification of the Aurignacian with the Szeletian or the Chatelperronian may document a period of co-habitation in the same regions of the makers of the former (anatomically modem humans) and the latter (Neanderthals). This suggests that either the Neanderthals could compete quite well with the anatomically modem humans, or that the latter had a relatively slow growth rate (or high death rate) and so took some time to totally swamp and replace the Neanderthals .

There appears to have been a gradual western expansion of the Aurignacian across Europe. Those who support the Replacement Hypothesis contend that the spread of the Aurignacian across Europe documents the actual movement of anatomically modem humans and that the Szeletian, Uluzzian, Chatelperronian and similar industries were manufactured by Neanderthals who were in contact with these modem peoples (Allsworth-Jones 1986a, 1986b; Harrold 1983, 1989; Stringer 1982, 1989b; Stringer et al. 1984; Stringer et al. 1981).

How and why anatomically modem humans replaced the Neanderthals are no doubt very complex matters .

As Gamble observed ...what we are seeing in the [European] archaeological record is a single change in human adaptive patterns which appears at different times in different areas; probably since it is related to the process of colonisation associated with anatomically modem Homo sapiens. (1986b:39)

As detailed above, Boaz et al. ( 1982) contend that anatomically modem humans migrated into western Asia during the warm parts of oxygen isotope stage 5, and that they migrated into Europe from there following the formation of a large land bridge across the Dardanelles at the boundary between oxygen isotope stages 2 and 3 which dates around 30,000 years BP (Table 5:1). They proposed that increasing desertification in central and northern Africa and parts of western Asia, and the shortage of resources it would have caused, formed the impetus that instigated the movement of anatomically modem humans into the cold but more productive steppe and tundra regions of Europe. Fogarty and Smith ( 1987) argued that this hypothesis is flawed because there is no convincing evidence for a prolonged period of severe desiccation in north Africa during this period, or that a land bridge formed at the Dardanelles.

Coexistence and Replacement The hominid sample (which appears to lack transitional specimens), and the lithic assemblages from Europe ( which document a westerly spread of the Aurignacian and the development of acculturated derivations of local Middle Palaeolithic industries) seem to support an external source for anatomically modem humans and therefore the Replacement Hypothesis. The metrical and non-metric analyses of cranial and mandibular data revealed a major separation between the Neanderthals and anatomically modem material (for example Figs. 5:7, 5:8, 5:9, 5:10, 5:11, 5:12, 5:13, 5:14, 5:15, 5:16, Tables 5:3, 5:4, 5:5). A major morphological discontinuity between European preNeanderthal and Neanderthal crania and western European anatomically modem crania was also revealed by the ordered similarity matrix analysis (Table 5:8).

There is however, increasing evidence to demonstrate that during the period from 60-30,000 years BP there were major climatic and temperature fluctuations across Europe (Gowlett 2001), which may have not only facilitated movement into Europe of anatomically modem humans from the south and east, but have caused periods of stress on the Neanderthal populations in Europe leading to episodes of both population contraction and expansion. These circumstances may have resulted in the abandonment of areas by Neanderthals and the subsequent repopulation by anatomically modem humans allowing a gradual expansion of the latter and a territorial contraction of the former.

Williams (1987) calculated evolutionary rate changes (using darwins and linear rate) from australopithecines through to anatomically modem humans. He found that the greatest amount of change was between Neanderthals and anatomically modem humans (Williams 1987), which again highlights the morphological difference between these two groups. Harvati (2003), analysing threedimensional coordinates of craniofacial landmarks using generalised procrutes analysis and multivariate statistical methods, found that there were greater intra-specific differences between Neanderthals and Late Pleistocene hominids and modem humans than was evident between

Mellars ( 1998b) has identified models to explain how prolonged periods of coexistence could occur in Europe. These include low population densities for both Neanderthals and anatomically modem humans, exploitation of different resources (discrete ecological or foraging niches) with limited overlap and successive rather 167

Skhul material, are more shelf -like with deeper ophryonic grooves , and more evenly thick, than those of the early European anatomically modem crania.

than simultaneous occupation of particular territories. In reality the situation was probably a combination of all of these models with different combinations in different areas. Mellars ( 1998b) goes on to propose two critical factors that led to the extinction and replacement of the Neanderthals - increasing population numbers of anatomically modem humans resulting in greater competition for resources , and, under these conditions , the inability of the Neanderthals to adapt to the rapidly deteriorating climatic conditions that occurred at 34,000 years BP (see also Stringer and Davies 2001).

The Neanderthals - Up Close and Personal If anatomically modem humans replaced the Neanderthals , was there any close contact between the two groups , especially considering that there appears to be a period of cohabitation within parts of Europe of some 10,000 years? Mitochondrial DNA (mtDNA) has been recovered from Neanderthal remains from opposite sides of the continent Feldho fer Cave, Germany, includin g the Neanderthal-type specimen (Krings et al. 1997 ; Schmitz et al. 2002) and hominid Vi-75 from Vindija Cave, Croatia (Krings et al. 2000)( also at Mezmaiskaya Cave in the northern Caucasus [Ovchinnikov et al. 2000]).

The anatomically modem human migrants may also have introduced new diseases , caused new selective pressures to be placed on the Neanderthal s by their presence, or could have had a higher growth rate . Zubrow ( 1986, 1989) has carried out demographic modelling that indicates that Neanderthal survival was impossible . He found for example , that if anatomically modem humans had a small demographic advantage over the Neanderthals of a 2% lower mortality rate there would be a rapid extinction of the Neanderthal s in approximatel y thirty generation s or one millennium (Zubrow 1989). Trinkaus ( 1986) has suggested (predominantly based on dental attrition) that anatomically modem humans had increased longevity compared with the Neanderthals , which would support Zubrow's ( 1989) demographic model.

The Feldhofer 1 sample produced a mtDNA sequence of 379 bas e pairs, which was compar ed with modem human sequenc es (Krings et al. 1997). The Feldhofer 1 sequence exhibited 27 differences from the modem reference sequence. The differences between Feldhofer I and the modem sample was three times greater than between pairs of modem humans, placing it outside the range of modem human variation. Adcock et al. (2001a) also found the Feldhofer 1 Neanderthal sequence to be separated from modem human sequences and also from ancient Australian sequences . A second individual from Feldhofer Cave has also provided a mtDNA sequence (Schmitz et al. 2002). Feldhofer 2 (NNl of Schmitz et al. 2002) produced a mtDNA sequence of 357 base pairs of the hypervariable region 1 (HVRl) that exhibited 23 differences from the modem reference sequence and one to four nucleotide differences from the other three Neanderthal sequences (Schmitz et al. 2002). Tree reconstructions placed the Feldhofer 2 mtDNA sequence with the other three Neanderthal sequences and separated them from the modem human sequences (Schmitz et al. 2002). Schmitz et al. (2002) also concluded that the mtDNA sequence differences between Feldhofer 1 and 2 (three differences) indicated that they were maternally unrelated.

Other possible explanations have been proposed (see Balter 2001 b), but what is clear is that anatomically modem humans replaced the Neanderthals in Europe (with or without a total replacement of gene pools). At present it is not possible to clearly identify from which morphological group the anatomically modem humans who colonised Europe derived. Brauer ( 1984a, 1984b, 1984c) for example, contends that Europe was colonised by anatomically modem humans from Africa and suggests marked similarities between the Omo 1 calvaria and Upper Palaeolithic European crania . The group that is most often seen as the ancestral stock from which the early European anatomically modem humans may have come are the hominids from Skhiil and Jebel Qafzeh. The revised dates for these hominids would support such a scenario (cf. Chapter 4). Skhiil 5 was often found to cluster with the Upper Palaeolithic crania (Fig. 2:3, 2:5, 2:7, 4:9, 4:11, 5: 16, 6:2, Table 2:2, 2:3, 2:4, 2:5, 4 :4, 4:5, 5:5). Jebel Qafzeh 9 was also found to cluster with European Upper Palaeolithic crania (Figs. 4:9, 4: 11, Table 4:4, 4:5). In an analysis of facial angles and indices , Stringer ( 1989b) found a close similarity between the Skhiil-Qafzeh sample and an Upper Palaeolithic European sample. Harvati (2003) also found that morphological distances were relatively close between Skhiil 5 and Jebel Qafzeh 9 and Upper Palaeolithic material.

Vi-75 provided a mtDNA sequence of 357 base pairs of the HVR-1 (hypervariable region) and 288 base pairs of HVR-2 (Krings et al. 2000). When compared with the Feldhofer 1 sequence, Vi-75 differed in 9 substitutions. The Feldhofer 1 and Vi-75 sequences differed to a modem reference sample by 34.9=/-2.4 substitutions and showed no closer relationship with a modem European reference sample than to reference samples from Africa or Asia (Krings et al. 2000), which would argue against the idea of genetic continuity between Neanderthals and early modem humans in Europe (Similar results were also obtained for the Mezmaiskaya sequence [Ovchinnikov et al. 2000]). On a schematic phylogenetic· gene tree the two Neanderthals grouped together and were separated from the modem samples (Krings et al. 2000) .

There are, however, morphological differences between these two groups. The western Asia hominids lack occipital "hemi-buns", which are common among the anatomically modem humans from Europe , although this feature is found on the Jebel Irhoud crania from north Africa. They also have flatter faces and less projecting nasal bones . In these features, the early anatomically modem European crania are more similar to the Neanderthals than they are to the Skhul and Jebel Qafzeh hominids. The large brow-ri dges, especially evident on the

The mtDNA diversity of the Feldhofer 1 and 2, Vi-75 and Mezmaiskaya sequences was found to be comparable to that of modem humans (Krings et al. 2000; Schmitz et al. 2002). 168

2001a, 2001b). The Neanderthal mtDNA sequences also seem to be distinct from sequences from anatomically modem ancient Australians of a similar or even older date (if the results are correct - see Chapter 6; Adcock et al. 2001a). Relethford (2001a) has argued that the results for Lake Mungo 3 obtained by Adcock et al. (2001a) demonstrate mtDNA lineage extinction, which could also explain the differences between Neanderthal and modem mtDNA sequences.

Ovchinnikov _et al. (2000) estimated that the age of the most recent common ancestor of the mtDNA of the eastern (Mezmaiskaya) and western (Feldhofer 1) Neanderthals was 151-352,000 years ago, which would coincide with the period during which the unique Neanderthal morphological pattern developed in Europe. The divergence of the Neanderthal and modem human mtDNA was estimated to be some 500-800,000 years ago, whereas the divergence of modem mtDNA sequences occurred sometime between 100-250,000 years ago (Krings et al. 1997; Ovchinnikov et al. 2000).

The mtDNA results would seem to exclude any significant interbreeding between Neanderthals and anatomically modern Upper Palaeolithic humans. As discussed by Caramelli et al. (2003), if Neanderthal mtDNA lineages are not found in modern reference samples because they disappeared due to random drift or stochastic lineage extinction, one could expect to find a closer link with mtDNA lineages from anatomically modem Upper Palaeolithic skeletons only some 5,000 years younger (than the Vindija Cave specimen), but this is not the situation. The conclusion that there was no significant interbreeding would be challenged if evidence of hybridisation could be identified within the fossil record, especially considering there is evidence that Neanderthals and anatomically modern humans were contemporaneous within regions of Europe. Skeletal remains from Portugal are proposed by some to be direct evidence of genetic contact between the two morphologically diverse groups.

Gutierrez et al. (2002) have suggested that the results of the analyses of the Neanderthal mtDNA can be questioned on methodological grounds. They concluded that the Neanderthal mtDNA sequences were not significantly different from and could not be considered an outgroup of modem humans. Gutierrez et al. (2002) also raised the possibility that the differences between Neanderthal and modem humans mtDNA may have been increased due to DNA damage. However, Gutierrez et al. (2002), like the other analyses, found the Neanderthal sequences were similar and clustered together and were different to that of the modem comparative samples. It is the level of difference that they questioned. The mtDNA HVRl has also been typed from two anatomically modem Upper Palaeolithic skeletons dated at around 24,000 years BP from Paglicci Cave, southern Italy (Caramelli et al. 2003). The mtDNA from these individuals, Paglicci-12 and Paglicci-25, is different to that of the Neanderthals and can be placed within the range of variation of modem humans (Caramelli et al. 2003). Caramelli et al. (2003) found Paglicci-12 to have 22-27 substitutions compared to Neanderthal sequences, whereas there was only a single substitution compared to the modem reference sample, while Paglicci-25 had 23-28 substitutions compared to Neanderthal sequences and only an average of 2.34 substitutions from the modem reference sample. On a multidimensional scaling analysis, the Paglicci sequences were placed within the range of variation of the modem reference sample and separated from the Neanderthal sequences (Caramelli et al. 2003). Also of interest is that the Paglicci-25 sequence was grouped into either the HV or pre-HV haplogroups, which are generally rare but are found in comparatively high frequencies among modern western Asian populations, while the Paglicci-12 sequence is grouped into the haplogroup N*, which is found in some western Asian populations (Caramelli et al. 2003). The results have been interpreted ·as indicating genetic continuity between anatomically modem Upper Palaeolithic and modern populations and a discontinuity with the Neanderthals (Caramelli et al. 2003). It should be noted that it remains difficult to totally discount possible contamination when dealing with DNA sequences so similar to those of modem humans (Abbott 2003; Barbujani and Bertorelle 2003; Cooper and Poinar 2000).

The site of Abrigo do Lagar Velho, has provided a burial containing a largely complete skeleton of a young child (Duarte et al. 1999). The skeleton and containing sediments were heavily stained with red ochre, possibly from an ochre-painted shroud, and there was an associated pierced shell, a practice similar to Gravettian burials from elsewhere in Britain and Europe (Duarte et al. 1999). The deposit overlying the burial has been dated to approximately 21,000 BP, while charcoal and faunal material directly associated with the burial have produced dates of 24,860+/-200 years BP (GrA-13310), 24,660+/260 years BP (OxA-8421) and 24,520+/-240 years BP (OxA-8423) (Duarte et al. 1999). These dates place the burial possibly 5000 years after the demise of the Neanderthals in the Iberian Peninsula and their replacement by anatomically modern humans (Mellars 1998a, 1998b). The skeleton is in a reasonably good condition although the skull and mandible were damaged by earth removal. The preserved left temporal bone is described as having a juxtamastoid eminence that extends to the same level as the mastoid process and being intermediate in size when compared to Neanderthal and early modem juveniles. The bony labyrinth of the inner ear is of the modern form and not the Neanderthal pattern (Bahn 2003; Hublin et al. 1996). The occipital is reported as displaying pitting similar to that seen on Neanderthals (Bahn 2003). The mandible is said to display a prominent development of a chin and a posteriorly retreating anterior symphysis with an angle close to a Neanderthal juvenile mean (Duarte et al. 1999). Dental dimensions of Lagar Velho 1 fall within the early modern human range (Duarte et al. 1999). The pubic length of Lagar Velho 1 also falls within the early modem human range, as does the curvature of the radius

These mtDNA sequence results have been interpreted as indicating that the Neanderthals did not contribute to the mtDNA pool of modem humans and therefore support the Replacement Hypothesis (Caramelli et al. 2003; Hoss 2000; Krings et al. 1997; Krings et al. 2000; Ovchinnikov et al. 2000; Ward and Stringer 1997; but see Relethford 169

The behavioural and biological interactions between Neanderthals and early modems during a period of cohabitation in Europe would have been complex in nature and may be difficult to recognise within the archaeological record. The skeletal and lithic data indicates that while there are some late sites, there was in general, a gradual replacement across Europe of the Neanderthals and Szeletian , Uluzzian, Chatelperronian and related industries by anatomically modem humans and the Aurignacian industry. As only anatomically modem humans have been associated with the Aurignacian it is probable that both phenomena are closely related and document the western migration of anatomically modem humans across Europe. There appears to be little conclusive evidence of in situ evolution from Neanderthals to anatomically modem humans in Europe with most "transitional" specimens problematic and mtDNA data precluding any significant interbreeding between Neanderthals and anatomically modem humans. Nordborg (1998) has concluded from statistical modelling that it is unlikely that interbreeding between Neanderthals and modem humans will be detected. Smith, however , contends that ... Neandertal genes may not have persisted to the present day, but those genes were there in the beginnings of modem European biological history. (2000:107) Such a contention will be difficult to assess either genetically or skeletally.

(Duarte et al. 1999). The tibia/femur length is closer to the Neanderthal ratio than that of early modems (Duarte et al. 1999). Lagar Velho 1 is also closer to Neanderthals than early modems in comparisons of femoral midshaft circumference to length (Duarte et al. 1999). Duarte et al. ( 1999) contend that Lagar Velho 1 had either very robust femora and tibia or shared the "hyperarctic" limb proportions with Neanderthals, the option they prefer. It has also been reported that Lagar V elho 1 has incisors showing a shovelling pattern typical of Neanderthals, and a suprainiac fossa (Balter 2001b). Duarte et al. ( 1999) conclude that Lagar Velho 1 presents a morphology that indicates an admixture between Neanderthals and anatomically modem humans and that the persistence of such a morphological mosaic into the Gravettian is indicative of significant hybridisation occurring throughout the Iberian Peninsula (see also Bahn 2003) . Whilst the limb proportions and the reported presence of a suprainiac fossa point towards the Neanderthals, there are no similar morphological patterns found at other Gravettian sites. Could this specimen be the only evidence of hybridisation in Europe or is Lagar V elho 1 just be a very stocky little boy? It is worth reiterating that the majority of specimens put forward as demonstrating hybridisation or evolutionary progression from Neanderthals to anatomically modem humans are either juvenile individuals or very fragmentary specimens.

It is interesting to note that with the chronological revisions that have occurred in Europe and western Asia and now place robust anatomically modem Homo sapiens earlier than the Neanderthals, some supporters of the Multiregional Hypothesis like Smith (1992) argue that any evidence of hybridisation excludes the possibility of "total replacement", whereas supporters of the Replacement Hypothesis such as Stringer and Brauer (1994) contend that hybridisation between the two groups cannot be equated with the Neanderthals evolving into anatomically modem Homo sapiens.

It is worth noting that Brauer observed .. .I see no hard evidence which would disagree with a replacement model in which the primary causes for the appearance of early modem humans were gene flow and population movements into Europe from outside. (1989: 139) In my eyes, a 'Hybridization and Replacement model' such as the 'AfroEuropean sapiens hypothesis' provides a reasonable explanation for the numerous differences between the Neanderthals and modem humans which were their direct successors, for the lack of clear transitional fossils, and for the evidence of some gene flow between the Neanderthals and the early modem Europeans. (1992:93)

The available evidence from Europe does not support any significant contribution of the Neanderthals to the biological or cultural history of anatomically modem humans and so is more supportive of the Replacement Hypothesis than the Multiregional Hypothesis .

170

CHAPTER SIX. AUSTRALASIA Introduction

Island Southeast Asia

Since the discoveries and pioneering work of Dubois, Weidenreich, and von Koenigswald, Australasia has been important in hominid evolutionary studies because of its numerous hominid remains. Java has produced the remains of many early hominids, whereas Australia has a large sample of late Pleistocene skeletal material (Table 6: 1). An assessment of the origin of anatomically modem humans in this region is difficult because of the lack of archaic Homo sapiens fossils, which necessitates comparisons between Homo erectus and anatomically modem Homo sapiens crania. Another problem is the lack of reliable dates and/or accurately established provenances for many of the hominid remains, although a basic chronology can be constructed. This is not to suggest that Australasia has not assumed a significant place within the debate surrounding the origin of anatomically modem humans, with the skeletal sample from the region forming a core component of the Multiregional Hypothesis (for example Frayer et al. 1993, 1994; Thome 1981a, 1984; Thome and Wilson 1977; Thome and Wolpoff 1981; Wolpoff 1980a, 1985, 1986a, 1989b; Wolpoff et al. 1984).

Southeast Asia has not produced any hominids that may conclusively be referred to as archaic Homo sapiens, although the taxonomic status of the Ngandong material continues to be debated (see Anton 1999; Hawks et al. 2000; Wolpoff et al. 2001). The area has produced a large Homo erectus sample and the remains of some potentially early anatomically modem Homo sapiens (Table 6: 1).

Indonesian Homo erectus The Indonesian island of Java has produced many early hominid remains (Fig. 6: 1). The exact stratigraphic provenance of most of the Javanese hominids is difficult to establish as many were either surface finds, recovered during construction work or purchased from collectors. There are also problems in the correlation of the localities of the few remains found in situ with the lithostratigrapical sequence at Sangiran (Pope and Cronin 1984; Sartono 1979; Sartono et al. 1981). The material is also from potentially reworked fluvial deposits, which complicates associations and interpretations of the hominid material, faunal collections and dating scenarios. Matsu'ura ( 1982,

Table 6:1. Australasian crania examined, abbreviations for Figures and Tables, and source of the data. Hominid Cohuna Cossack Keilor KowSwamp 1 KowSwamp 5 Kow Swamp 15 Lake Mungo 1 Lake Mungo 2 Lake Mungo 3 Lake Nitchie Lake Tandou Mossgiel Niah Cave Ngandong 1 Ngandong 5 Ngandong 6 Ngandong 10 Ngandong 11 N gandong Mean Pithecanthropus erectus 1 [Trinil 2] Pithecanthropus erectus 2 [Sangiran 2] Sangiran 17 Tabon Cave Talgai Wajak 1 Wajak2 Willandra Lakes Hominid 50

Orieinal Or Cast Cast Cast and Original Cast Cast and Original Cast and Original Cast Cast and Original Published Cast and Original Cast Original Cast Cast Published Published Published Published Published Published Published and Cast Published and Cast Cast and Published Cast and Published Cast Cast Cast Cast and Original

Abbreviations

---

---------

---Nl NS N6 NlO Nll Nm Pel Pe2 Sl7

--

---

WI

--

Principal Source Of Data Cast Cast and Original Cast Cast, Thome 1976 Cast, Thome 1976 Cast Cast, Thome 1971a Bowler et al. 1970 Cast Cast Original Cast Cast Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Weidenreich 1943b Cast, Thome and Wolpoff 1981 Cast, Baker 1978, Macintosh 1978 Cast Cast Cast Cast and Original

Note that for the Ngandong material the traditional numbering system has been followed, although the site name Ngandong has been substituted for Solo (cf. Santa Luca 1980:16). 171

PHILIPPINES

SUNDA

Ngandong

Sangiran

17

iP

Wajak

1

Figure 6:1. Map showing Southeast Asian sites mentioned in the text.

1986) attempted to establish the stratigraphical provenances and relative dating of some of the Sangiran hominids by conducting fluorine studies of the fauna, including hominid remains, from the site. Matsu'ura concluded that Sangiran 3, 12 and 17 derived from the Kabuh Fonnation between the lower and middl e tuff, and

that Sangiran 1b, 2, 4 and 6 derived from the Grenzbank basal conglomerate of the Kabuh Formation , while Sangiran 5 may have come from this same horizon or from the upper part of the Pucangan Formation . This study did demonstrate that the hominid material was distributed throughout the deposits.

172

Matsu'ura ( 1982, 1986) obtained a "possible" fluorine age of 1.3 m.y.a. for Sangiran 5. The other ages for Javanese hominids obtained by Matsu'ura range between 0.8 to 1.1 m.y.a. Siesser and Orchiston (1978) assigned a minimum age of 1.6 m.y.a. for Pithecanthropus Mandible CSangiran 9, a surface find, on the basis of the foraminiferal species present in the claystone adhering to it, but as Pope and Cronin (1984:379) point out, this study was "based on an inadequate and obviously reworked sample with highly debatable stratigraphic ranges" (cf. also Watanabe and Kadar 1985).

There are also major problems with the dating of the various deposits and formations in Java, and as mentioned above, cross-correlation between localities is difficult. For these reasons only a broad chronological framework for Javanese hominid material can be established. Detailed discussions of the problems of the geological and temporal framework of the Indonesian hominid finds can be found elsewhere and so only a resume of the various debates will be provided here (cf. de Vos 1994; Groves 1985; Huffman, 2001; Langbroek and Roebroeks 2000; Matsu'ura 1986; Orchiston and Siesser 1982; Pope 1982 1983 1985; Pope and Cronin 1984; Sartono 1983/84, 1985; Semah 1982, 1984, 1986; Semah et al. 1981; Semah et al. 2000; Shutler and Braches 1985; Swisher et al. 1994; Watanabe and Kadar 1985). It will suffice to quote Pope who stated Estimates of the age of the early Javanese hominids were originally based in ill-defined lithostratigraphic ...and overly simplistic biostratigraphic ...sub-divisions that failed to recognize the complex nature of Javanese stratigraphy. ( 1983 :4988)

Swisher et al. (1994) have obtained weighted mean ages of l.81+/-0.04 m.y.a. for a site at Perning near Mojokerto, where a juvenile calvaria may have been recovered and I.66+/-0.04 m.y.a. for the Sangiran location (from where Sangiran 27 and 31 were recovered) from argon laserincremental heating analyses of hornblende separated from volcanic pumice. These dates are significantly older than other proposed dates for Javanese Homo erectus hominids and are comparable to the earliest dates for Homo erectus hominids in Africa and Europe. Whilst the dates obtained by Swisher et al. ( 1994) appear to be technically correct, their direct association with the hominids from the two sites remains open to question. It is also worth noting that two possible find sites for the Mojokerto calvaria have been identified (Huffman 2001 ).

Von Koenigswald initially proposed a faunal sequence for Java including the Lower Pleistocene (D)Jetis Fauna (Pucangan Formation), the Middle Pleistocene Trinil Fauna (Kabuh Formation) and the Upper Pleistocene Ngandong Fauna (Notopuro Formation), which covers the hominid bearing deposits. However, this succession of faunas did not occur at any single locality. Reinterpretation of the faunal material from Java has been undertaken and resulted in the suggestions that the Jetis Fauna may actually be younger than the Trinil Fauna (Leinders et al. 1985; Sartono 1983/84, 1985; Sondaar 1984; de Vos 1985; de Vos and Sondaar 1982; de Vos et al. 1982), and that these two faunal assemblages share many "guide fossils" that were initially used to differentiate them (Matsu'ura 1982; Sartono 1983/84; Sartono et al. 1981; Watanabe and Kadar 1985). Bartstra (1983) and Hooijer (1983), however, contend that the Dubois Collection is unsuitable for biostratigraphical studies such as those done by de Vos and others, because it is a mixed collection of fossils from Lower and Middle Pleistocene, and possibly Lower Pliocene, deposits. They concluded that von Koenigswald's faunal sequence was essentially correct, and that the Jetis Fauna was older than the Trinil Fauna. Groves (1985) also argued that von Koenigswald's sequence was essentially correct, citing as evidence the presence of the remains of a more archaic tiger from Kedungbrubus than from Trinil. Further clarification is still needed.

Based on lithologic and paleogeographic evidence Huffman (2001) supported the date of 1.81+/-0.04 m.y.a. obtained by Swisher et al. (1994) for the Mojokerto site and by association the Mojokerto calvaria. De Vos and Sondaar ( 1994) questioned the geological context of the dated samples which they contend may have been reworked and redeposited and the magnetostratigraphic interrelations proposed by Swisher et al. (1994) including the attribution of the normal polarity to the Olduvai event and not the younger Jarmillo event (see also Hyodo et al. 2002). Swisher (1994) has reaffirmed the relationship of the dates to the hominid sites and dismisses the magnetostratigraphic interpretations of de Vos and Sondaar (1994). Semah et al. (2000) suggested that Swisher et al. ( 1994) most probably dated the underlying "lower lahar" deposit and not the fossil-bearing Pucangan Formation. Semah et al. (2000) undertook palaeomagnetic studies and argon dating of the volcanic "lower lahar" layer at the base of the fossil-bearing deposits at Sangiran. They concluded that the maximum theoretical age for the arrival of hominids at Sangiran was 1.7 m.y.a., although there appeared to be gap prior to the sedimentation of the fossilbearing Pucangan Formation (Semah et al. 2000).

Absolute dating of the early Javanese hominid localities remains problematical and confused, with varying methods and studies providing different results. For instance, Orchiston and Siesser (1982) placed the Kabuh-Pucangan boundary between 1.2 and 1.4 million years ago (m.y.a.) based, predominantly, on an average date of 0.83 m.y.a. for four pumice samples from the lower Kabuh (Bapang) Formation at Sangiran, and a 1.9 m.y.a. date from the upper levels of the Pucangan Formation, whereas palaeomagnetic studies (Semah 1982; Semah et al. 1981) indicate that the Bruhnes/Matuyama boundary lies near the Grenzbank basal conglomerate of the Kabuh Formation.

Langbroek and Roebroeks (2000) have argued that the silicate glass tektite layer between the Lower and Upper Tuffs of the Kabuh Formation, which overlies the Pucangan Formation, demonstrates that the middle Kabuh Formation and the associated hominid remains cannot be older than 800,000 years BP. This conclusion is based on radiometric dating of tektites and the stratigraphical position of tektites in deep-sea cores. Langbroek and Roebroeks (2000) concluded that all of the Sangiran 173

Santa Luca (1980), however, has provided taphonomic evidence to suggest that the Ngandong Fauna is a mixed assemblage with the non-hominid fauna representing a death-assemblage buried after minimal exposure, while the hominid remains appear to have been redeposited into younger levels. This suggestion is consistent with the arrangement of the hominid remains within the terrace. The non-hominid fauna is well preserved with complete and articulated vertebral columns and crania complete with associated mandibles, whereas the hominid material is fragmented and consists predominantly of calvaria. The hominid sample is composed of only the most durable structures, with the calvaria displaying evidence of surface damage and lacking the facial skeleton, which could be the result of rolling and transportation of the crania by water (Boaz and Behrensmeyer 1976). This pattern is compatible with the suggestion that the hominid sample had been redeposited. Coon (1962:390) , however , stated that the hominid remains "had not been moved or rolled". A recent examination has supported Santa Luca 's (1980) contention . Following a limited taphonomic analysis of hominid (Ngandong 9 and 10) and faunal material, Westaway (2002) observed that the hominids and fauna had distinct and different taphonomic alterations that indicated they had been deposited in at least two different hydrological events. Westaway (2002) even suggested that Ngandong 10 could have had multiple episodes of transportation and burial. The degree of mineralisation, lack of obvious weathering and the presence of pressure distortion on the hominid material prompted Westaway (2002) to infer that the Ngandong hominids were older than the faunal material. The idea that the Ngandong hominids were redeposited into the younger Solo High Terrace has not been universally accepted (see Bartstra and Bosoeki 1989; Bartstra et al. 1988; Swisher et al. 1996). Bartstra and Bosoeki dismissed the contention by observing that .. .for anyone who is familiar with the field situation around Ngandong , the question is where the hypothetical parent sediments of the N gandong skulls might be situated. (1989:243) and that It can hardly be assumed that 11 skulls (and 2 tibia) were transported together by stream action along a meandering river course for more than 30 km. (1989:243) Westaway ' s (2002) analysis, although limited in the material studied, indicated that Ngandong 10, at least, could have had multiple episodes of transportation and burial and so the transportation of the hominid material over the 30km identified by Bartstra and Bosoeki could have occurred in a number steps and over a prolonged period.

hominid remains can be placed between the onset of the Jaramillo subchron and the Bruhnes/Matuyama transition, that is between 1.07 m.y.a. and 780,000 years BP. Barstra and Basoeki (1989) have also argued that the Kabuh Formation is essentially congruent with the Bruhnes normal epoch and dates to between 715,000 year BP and 250,000 years BP (Barstra and Basoeki 1989:Fig. 3). Research on the Indonesian island of Flores suggests that stone tool making hominids were not present on the island 900,000 years ago, but they had arrived by 840,000 years BP (Morwood et al. 1998; Morwood et al. 1999). The appearance of stone tool making hominids on Flores also coincides with a faunal change from an endemic fauna including pygmy stegodons and giant tortoises to a faunal sample including continental southeast Asian elements such as large stegodons , which has prompted the suggestion that the hominids caused the extinction of the endemic fauna (Gibbons 1998). It is worth noting that some questions remain about the identification of the stones as tools (Gibbons 1998) . The other significant issue with this research is that even at times of lowest sea levels water crossings were necessary to reach Flores, and so the archaeological evidence has been interpreted as indicating that the Flores stone tool making hominids (most probably Homo erectus) were capable of constructing and using water craft (Morwood et al. 1998; Morwood et al. 1999). An age for the early Javanese Homo erectus hominids of less than 1.3 m.y.a. would fit with an estimate of 1.25 m.y.a . for a period of maximum exposure of the Sunda shelf, the combined land mass of mainland and island southeast Asia (Berggren and van Couvering 1979), which would facilitate the movement of hominids, and other fauna, into the region (Leinders et al. 1985). Reviews of the various dating schemes and additional references may be found in the publications listed above. The available radiometric , palaeomagnetic , and biostratigraphical evidence suggests that the earliest Indonesian hominids are younger than 1.3 m.y.a., with most probably dating to between 500,000 years BP and 800,000 years BP. It is interesting to note that computer simulations of hominid dispersal from Africa have hominids arriving in Indonesia after 1.5 m.y.a. with a mean date of 0.97 m.y.a. from 30 default simulation runs (Mithen and Reed 2002) . The date of the N gandong hominid sample remains difficult to ascertain with any certainty . The hominids were recovered from the 20-metre terrace (referred to as the Upper or High Terrace) of the Solo River near Ngandong, central Java. (Fig. 6: 1; Santa Luca 1980; Oakley et al. 1975; Weidenr eich 1951). They were not localised in any particular spot or within a single layer, but were irregularly distributed throughout the entire site, and so while they can be regarded as a sample they do not necessarily represent a single biological population. The Solo High Terrace, which is referred to the Notopuro Formation , contained abundant, predominantly extant, mammalian fauna (the Ngandong Fauna), and so is usually considered to be of Upper Pleistocene age (Jacob 1978; Santa Luca 1980; Sartono 1983/84, 1985). This has meant that some scholars, such as Coon ( 1962), have given the Ngandong hominids an Upper Pleistocene date.

Bellwood (1997) suggested that the Ngandong hominid sample may be the remains of carnivore kills, whereas Wolpoff (1986c) proposed that the sample displayed evidence of hominid behaviour . Both use evidence from the South African australopithecine sites to support their proposal s. Jacob ( 1972) dismissed the suggestion (cf. Coon 1962; Weidenreich 1951) that any of the early Indonesian hominids, including the Ngandong sample, demonstrated head-hunting and brain-eating. 174

Bellwood ( 1997) outlines the ranges of dates that have been proposed for the Ngandong material. Lestrel (1975) quoted an unpublished potassium-argon date of 310,000+/l 00% years BP for the Solo provenience, but the large standard error makes this date of no use. Lestrel and Read (1973), using regression estimates of brain size on time, predicted that the N gandong hominids were dated between l 96,000