The Human Fossil Record, Craniodental Morphology of Early Hominids (Genera Australopithecus, Paranthropus, Orrorin), and Overview (Volume 4)

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THE HUMAN FOSSIL RECORD Series Editors JEFFREY H. SCHWARTZ Department of Anthropology University of Pittsburgh Pittsburgh, Pennsylvania IAN TATTERSALL Department of Anthropology American Museum of Natural History New York, New York

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THE HUMAN FOSSIL RECORD Volume Four Craniodental Morphology of Early Hominids (Genera Australopithecus, Paranthropus, Orrorin) and Overview

Jeffrey H. Schwartz Department of Anthropology University of Pittsburgh Pittsburgh, Pennsylvania

Ian Tattersall Department of Anthropology American Museum of Natural History New York, New York

(\|/)WILEY-LISS A JOHN WILEY &, SONS, INC., PUBLICATION

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This book is printed on acid-free paper. © Copyright © 2005, text by Jeffrey 11. Schwartz and Ian Tatlersall; illustrations and photographs by Jeffrey 11. Schwartz (with exception* a» noted in the Preface). All rights reserved. Published by John Wiley &, Sons, Inc., I loboken, New Jersey, Published simultaneously in Canada. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted under Section 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of the Publisher, or authorization through payment of the appropriate per-copy fee to (he Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, 978-750-8-100, ^x 978-6468600, or on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the Permissions Department,John Wiley ck.Sons, Inc., I l l River Street, I loboken, NJ 07030, (201) 7486011, fix (201) 74S-6008. Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives or written sales materials. The advice and strategies contained herein may not be suitable for your situation. You should consult with a professional where appropriate. Neither the publisher nor author shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. The publisher and the author make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of fitness for a particular purpose. This work is sold with the understanding that the publisher is not engaged in rendering professional services. The advice and strategics contained herein may not be suitable for every situation. In view of ongoing research, equipment modifications, changes in governmental regulations, and the constant flow of information relating to the use of experimental reagents, equipment, and devices, the reader is urged to review and evaluate the information provided in the package insert or instructions for each chemical, piece of equipment, reagent, or device for, among other things, any changes in the instructions or indication of usage and for added warnings and precautions. The fact that an organization or Website is referred to in this work as a citation and/or a potential source of further information does not mean that the author or the publisher endorses the information the organization or Website may provide or recommendations it may make. Further, readers should be aware that Internet Websites listed in this work may have changed or disappeared between when this work was written and when it is read. No warranty may be created or extended by any promotional statements for this work. Neither the publisher nor the author shall be liable for any damages arising herefrom. For general information on our other products and services please contact our Customer Care Department within the U.S. at 877-762-2974, outside the U.S. at 317-572-3993 or fax 317-572-4002. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print, however, may not be available in electronic format. Library of Congress Cataloging-in-Publication Data is available. 0-471-31929-5

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Preface

CONTENTS

ix

Lothagam Lukeino Maka Makapansgat Male ma Olduvai Gorge Omo Valley, Lower (Shungura, Usno) Peninj (Lake Natron) Stcrkfontein Swartkrans Tabarin (Tugen Hills) Taung (Taungs) Turkana, West (Lomckwi, Lokalalei, Nachukui)

PARTI. INTRODUCTION Descriptive Protocol Descriptive Format Anatomical Terminology Figures Abbreviations Maps Layout of En tries

3 3 4 13 14 18

PART 2. SITE ENTRIES Allia Bay Belohdclie Chcsowanja Drimolen Fejej Hadar Kanapoi Koobi Fora (East Turkana, East Rudolf) Kromdraai Laetoli (Laetolil, Garusi)

23 30 33 37 48 50 118

200 202 209 219 243 245 254 294 298 374 437 439 448

PART 3. HOMINID CRANIODENTAL MORPHOLOGIES: A N OVERVIEW

Introduction Systematic Approach to the Hominid Fossil Record The Family Hominidae and the Earliest Ho mini ds

141 166 182

vii

465 466 467

CONTENTS

VIII

Operational Problems in the Alpha Taxonomy of the "Early Hominids"

471

Thc'Australopiths" Southern Africa Eastern Africa: The "Robusts" Other Australopiths" Australopithecus anam ens is Australopithecus afarensis "Early Homo"

473 473 479 482 482 483 486

The Ubiquitous Homo erectus: Species or

490

Grab-Bag? Homo erectus and Its Putative Relatives 490 in Java Putative Homo erectus in China 493 Putative Homo erectus from Africa 494

Putative Homo erectus from Europe

493

Middle and Late Pleistocene Hominids of Europe

500

Early Middle Pleistocene Hominids Homo heidelbergensis and Its Putative Relatives

500 502

T h e Neanderthals and Related Forms

504

Homo sapiens and "Archaic Homo sapiens" 506 Homo sapiens and Suggested 507 Close Relatives Other Members of the "Archaic 508 Homo sapiens" Group from the Levant and Africa Coda Appendix

510 552

PREFACE

This is the fourth volume in a series dedicated to the standardized description and illustration of the principal fossil specimens that document the hominid biological past. O u r hope is that these volumes will provide a comparative resource that may be used by those many interested individuals who lack the time and/or resources to examine first-hand the swelline number of original fossils that document the long story ot hominid evolution. The first two volumes, authored by us, are devoted to skulls and dentitions that have been allocated to the genus Homo (Volume 1 covers Europe, and Volume 2 Africa and Asia); the third volume, by Ralph Holloway, Doug Broadfield and Michael Yuan, discusses cranial endocasts; and in this tourth volume we describe African craniodental remains that have been attributed to the genera Australopithecus^ Paranthropas and Orrorin.

possible to sort out the species-level variety of hominids to general satisfaction, it will be premature to proceed to higher-level analyses. We do, however, provide at the end of this fourth volume a general survey of the morphologies we have encountered in the compilation of our three craniodental volumes, and we note some of the implications of those morphologies. Another regret is the absence from this resource of several early hominid or putatively hominid genera such as the recendy named Sahelanthropns> Kenyanthropus and Ardipithecits\ but we hope to be able to include them in future contributions to this series. We begin this volume with an abbreviated account of our descriptive protocol (for a more comprehensive treatment of this subject, the reader is referred to Volume 1 of this series). As in earlier volumes, the entries are organized alphabetically by site; and to the morphological descriptions are added details ot discover}*, dating, archaeological context and so forth. Description remains paramount, however. As before, where morphological variety seems to dictate we have grouped fossils from a single site into various morphs. In this volume, particularly, many of these morph groupings will be unfamiliar, and many are necessarily tentative. We have thus been careful to describe as many individual fossils as possible, and the reader should be able to locate these individual descriptions

As our work on the series progressed, it became clear that we would have to stop short of our original goal ot concluding this fourth volume with a general survey of hominid systematics and phylogenetic relationships. For although we fully recognize that not all morphs represent their own species, the diversity of morphologies we unearthed in the process of making detailed descriptions of so many fossils makes it clear that it is too soon to claim even that a rational alpha taxonomy of fossil hominids is in sight. Until it is IX

X

PUKFACE

readily, even where the order of listing is perhaps unexpected. Clearly, the definitive account of the human fossil record will never be written. Partly this is for the best of reasons: the paleoanthropological record is already very extensive, and it is growing at a rate with which it is hard to keep up. Partly, though, it reflects the fact that certain human fossils, even ones that have been comprehensively published, arc surrounded by a wall of protectionism that constitutes a major stumbling block in what is after all a comparative science. Thus, for example, the type materials of two proposed species of Australopithecus, named and described in leading scientific journals several years ago, remain at this writing off-limits to the general paleoanthropological community (and absent from this volume) due to their describcrs* steadfast resistance to any independent verification. These are the species Australopithecus garhi and Australopithecus bahrelghazali, and they are joined by described specimens (e.g., the Konso cranium) that are alleged to extend significantly the morphological ranges of their species (in that particular case, Paranthropus boisei), but that are likewise unavailable for general examination. T h e prevailing problems of access to published type materials run directly counter not only to the spirit but to the letter of the International Code of Zoological Nomenclature, in which (4th edition, 1999) Paragraph 72.10 states that "Holotypes, syntypes, lectotypes and neotypes . . . . are to be held in trust for science by the persons responsible for their safe-keeping (p. 79)". T h e associated Recommendation 72F adds that "Every institution in which name-bearing types are deposited should . . . make them accessible for study" (p. 79). 1 his stricture applies, of course, not only to institutions but also to those who influence their policies. Until access to published type and associated specim e n h permitted to researchers outside the closed dec-aibing cliques, the species based on them must perforce be regarded by the majority of us as hypothetical constructs, equivalent to such entities as Hseckel*s "Pithecanthropus alalus." These remarks having reluctantly been made, it is gratifying to acknowledge the extraordinary assistance and hospitality extended to us by the great majority of those responsible for the care of the fossils that make up the human biological record. Without the active help of many colleagues this project would never have been possible, and warm welcomes and exceptional kindness all over the world have transformed a potentially

Herculean labor into a pleasure. For this fourth volum we extend our warmest thanks and appreciation lto ti ° u r~n : i._ ... ^ .. tile following, who gave us access to fossils °r provided other valuable help: Tim Bromagc, Ron Clark Nicholas Conard, Eric Delson, Heidi Fouric, Eustace Gitonga, Miriam Haidlc, Andrew Hill, p. Clark Howell, Ato Jara Hailc Mariam, Yusuf Juwaycyi Donatius Kamamba, Teresa Kearney, Kebedc Workc' Andre Keyscr, Bill ICimbel, Beverly Kramer, Kathy Kuman, Kevin Kuykcndall, Mcavc Leakey, Nasser Richard MalitJ o h n Maringah, Mamitu Yilma, Michael Mbago, Emma Mbua, Jacopo Moggi-Cecchi, C. S. Msuya', Charles Musiba, H. M. Nguli, Martin Pickford', Stcphany Potze, Yocl Rak, Issa Ramadhan, Fricdcmann Schrcnk, Horst Scidler, Brigittc Scnut, Francis Thackeray, Phillip Tobias, Alan Walker and Tim White (Maka and Bclohdelic). Wc would also like to take this opportunity to thank Harry Widianto of the Archaeological Service in Yogyakarta, and Mr. Himawan of the Museum Mpu Tantular in Surabaya, for access to Sangiran and Ngawi fossils described in Volume 2. The institutions in which the fossils described here reside, and whose official cooperation was obviously essential, are listed individually by site entry. Wc arc more than grateful to all of them, and thank them for permission to publish photographs of specimens in their care. Photographs of fossils from the Kenya National Museums and the National Museum of Ethiopia arc copyright of those institutions, and the image of Gran Dolina ATD6-5 is © Javier Trucba. Ken Mowbray kindly took the image of the Zhoukoudian reconstruction. All other photographs arc copyright © Jeffrey H . Schwartz. We thank Tim D. Smith for the black-and-white drawings, and Ken Mowbray for the maps. Many other friends and colleagues have also been indispensable in making these volumes a reality. Our initial editor at Wiley, Robert Harington, enthusiastically embraced the notion of this scries, which could not have come to fruition without the commitment or Luna Han and Thorn Moore, who steered it through to completion. Also at Wiley, numerous individuals at all stages of production and marketing deserve our warmest appreciation, particularly Kristin Cooke Fasano, who oversaw production. At the American Museum of Natural I listory Ken Mowbray and Shara Bailey rendered valuable help. At the University o( Pittsburgh, Michelle Ray, first at Photographic Services, and later independently, undertook the painstaking task of scanning all the black-and-white negatives and enhancing each image to bring out a>

PRE 'ACE

Xi

%

much detail as possible. She also compiled the comparative plates in the concluding discussion, with exacting attention to detail. To all of these individuals we offer our warmest thanks. In addition to our personal financial contributions, support for this work was provided by funds administered through the Department of Anthropology of the American Museum of Natural History, as well as by grants from the L. S. B. Leakey Foundation, John Wiley & Sons, Ncvraumont Publishing Co., and, at the University of Pittsburgh, the Central Research Development Fund, the University Center for International Study, and the Nationality Rooms Programs (J. G. Bowman). Finally, it should be noted that no palcoanthropologists embarking on a project such as this one could ever ignore the fact that they are standing on the shoulders of some very illustrious predecessors. Notable among these forerunners are the authors and editors of the Catalogue des Hommes Fossiles,

edited by H. V. Vallois and H. L. Movius and published in 1953; the three volumes of the Catalogue of Fossil Hominids, edited by K. P. Oakley, B. G. Campbell and T. I. Molleson, and published and revised between 1967 and 1977; and M. H. Day's multicdition Guide to Fossil Man, which first appeared in 1965. New information relevant to these catalogues is available via the invaluable series Horn in id Remains: An Up-Date, edited on an ongoing basis by R. Orban, P. Semal and colleagues. None of these works has or had exactly the same intentions as this one; for example, the Catalogue of Fossil Hominids aimed at comprehensiveness of sites but ignored morphology and illustration, whereas the Guide to Fossil Man did provide some general morphological information and illustration but was more selective in site choice. Nevertheless, we are keenly aware that we are following a road that has already been partly trodden, and that our task has thereby been rendered easier.

PART

ONE

INTRODUCTION

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