New Interpretations of Ape and Human Ancestry

ADVANCES IN PRIMATOLOGY

Series Editors:

W. PATRICK LUCKETT University of Puerto Rico Sanjuan, Puerto Rico

CHARLES R. NOBACK Columbia University New York, New York

Editorial Board:

RUSSELL L. CIOCHON, University of California, Berkeley, California

JOHN F. EISENBERG, Florida State Museum, Gainesville, Florida

MORRIS GOODMAN, Wayne State University School of Medicine, Detroit, Michigan

F. A.JENKINS,Jr., Harvard University, Cambridge, Massachusetts

FREDERICK S. SZALAY, Hunter College, New York, New York

TIlE PRIMATE BRAIN Edited by Charles R. Noback and William Montagna

MOLECULAR ANTHROPOLOGY: Genes and Proteins in the Evolutionary Ascent of the Primates Edited by Morris Goodman and Richard E. Tashian

SENSORY SYSTEMS OF PRIMATES Edited by Charles R. Noback

NURSERY CARE OF NONHUMAN PRIMATES Edited by Gerald C. Ruppenthal

COMPARATIVE BIOLOGY AND EVOLUTIONARY RELATIONSHIPS OF TREE SHREWS Edited by W. Patrick Luckett

EVOLUTIONARY BIOLOGY OF THE NEW WORLD MONKEYS AND CONTINENTAL DRIFT Edited by Russell L. Ciochon and A. Brunetto Chiarelli

NEW INTERPRETATIONS OF APE AND HUMAN ANCESTRY Edited by Russell L. Ciochon and Robert S. Corruccini

A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher.

New Interpretations of Ape and Human Ancestry

Edited by

RUSSELL L. CIOCHON University l!! California Berkeley, California

and

ROBERT S. CORRUCCINI Southern Illinois University Carbondale, Illinois

PLENUM PRESS • NEW YORK AND LONDON

Library of Congress Cataloging in Publication Data

Main entry under title:

New interpretations of ape and human ancestry.

(Advances in primlttology) Includes bibliographical references and indexes. 1. Primates, Fossil-Addresses, essays, lectures. 2. Human evolution-Addresses,

essays, lectures. I. Ciochon, Russell L. II. Corruccini, Robert S. III. Series. QE882.P7N48 1983 569'.8 83-2175 ISBN 978-1-4684-8856-2 ISBN 978-1-4684-8854-8 (eBook) DOI 10.1007/978-1-4684-8854-8

© 1983 Plenum Press, New York Softcovcr reprint of the hardcover I st edition 1983 A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013

All rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

Dedicated To

SHERRY WASHBURN Whose Ideas Are Embodied

In Many Chapters Of This Volume

and

CLARK HOWELL In Recognition Of His Considerable Contributions To Paleoanthropology

In Memory of G. H . R. VON KOENIGSWALD

A legendary figure in the field of paleoanthropology

G . H. R. von Koenigswald 1902-1982

Contributors

Peter]. Andrews Department of Paleontology British Museum (Natural History) Cromwell Road London SW7 5BD, England

Marietta L. Baba Department of Anthropology College of Liberal Arts Wayne State University Detroit, Michigan 48201

Raymond L. Bernor College of Medicine Department of Anatomy Laboratory of Paleobiology Howard University Washington, D.C. 20059

Noel T. Boaz Department of Anthropology New York University New York, New York 10003

s. R. K. Chopra Department of Anthropology Panjab University Chandigarh-160014, India

Russell L. Ciochon Department of Paleontology

(temporary research appointment) University of California Berkeley, California 94720

Robert S. Comu:cini Department of Anthropology Southern Illinois University Carbondale, Illinois 62901

ix

John E. Cronin Departments of Anthropology and

Biology Peabody Museum Harvard University Cambridge, Massachusetts 02138

Linda L. Darga Department of Anatomy School of Medicine Wayne State University Detroit, Michigan 48201

Louis de Bonis Laboratoire de Paleontologie des Ver-

t'ebres et Paleontologie Humaine Universite de Poitiers Faculte des Sciences 86022 Poitiers, France

Richard Dehm Institut fUr Palaontologie und histo­

rische Geologie Universitat Munchen 8000 Munchen 2, West Germany

Dean Falk Department of Anatomy and Caribbean

Primate Research Center University of Puerto Rico Medical Science Campus San jqan, Puerto Rico 00936

John G. Fleagle Department of Anatomical Sciences Health Sciences Center State University of New York Stony Brook, New York 11794

X CONTRIBUTORS

David G. Gantt Institute of Dental Research University of Alabama School of

Dentistry University Station Birmingham, Alabama 35294

Morris Goodman Department of Anatomy School of Medicine Wayne State University Detroit, Michigan 4820 I

Leonard O. Greenfield Department of Anthropology Temple University Philadelphia, Pennsylvania 19122

Donald C. Johanson Institute of Human Origins 2700 Bancroft Way Berkeley, California 94704 and Laboratory of Physical

Anthropology Cleveland Museum of Natural History Cleveland, Ohio 44106

Richard F. Kay Department of Anatomy Duke University Medical Center Durham, North Carolina 27710

William H. Kimbel Laboratory of Physical Anthropology Cleveland Museum of Natural History Cleveland, Ohio 44106 and Departments of Anthropology and

Biology Kent State University Kent, Ohio 44240

Arnold G. Kluge Museum of Zoology and Department of

Ecology and Evolutionary Biology The University of Michigan Ann Arbor, Michigan 48109

Adriaan Kortlandt Vakgroep Psychologie en Ethologie der

Dieren Universiteit van Amsterdam Nieuwe Achtergracht 127 IOIH WS Amsterdam, The Netherlands

Jerold M. Lowenstein Department of Medicine (122 MR2) University of California San Francisco, California 94143

Larry L. Mai Division of Orthopaedic Surgery UCLA School of Medicine and Department of Anthropology University of California Los Angeles, California 90024

Henry M. McHenry Department of Anthropology University of California Davis, California 95616

Mary Ellen Morbeck Department of Anthropology University of Arizona Tucson, Arizona 85721

Martin Pickford National Museum of Kenya P.O. Box 40658 Nairobi, Kenya

David R. Pilbeam Department of Anthropology Harvard University Cambridge, Massachusetts 02138

K. N. Prasad Geological Survey of India 4-3-542, Bogulkunta Tilak Road, Alladin Building H yderabad-50000 I, India

Michael D. Rose Department of Anatomy New Jersey Medical School University of Medicine and Dentis-

try of New Jersey 100 Bergen St. Newark, New Jersey 07103

Vincent M. Sarich Departments of Anthropology and

Biochemistry University of California Berkeley, California 94720

Elwyn L. Simons Duke University Center for the Study

of Primate Biology and History Durham, North Carolina 27705

C. H. R. von Koenigswald Section of Paleoanthropology Senckenberg Museum 6000 Frankfurt I, West Germany

Alan C. Walker Department of Cell Biology and

Anatomy The Johns Hopkins University School of Medicine Baltimore, Maryland 21205

Steven C. Ward Department of Anthropology Kent State University Kent, Ohio 44242 and Department of Anatomy Northeast Ohio Universities College of

Medicine Rootstown, Ohio 44272

CONTRIBUTORS xi

Tim D. White Department of Anthropology University of California Berkeley, California 94720

Milford H. Wolpoff Department of Anthropology University of Michigan Ann Arbor, Michigan 48109

Adrienne L. Zihlman Department of Anthropology University of California Santa Cruz, California 95064

Preface

In the field of paleoanthropology there is perhaps no more compelling issue than the origin of the human lineage. From the 19th-Century proclamations of Darwin, Huxley, and Haeckel up to the present-day announcements of authorities such as Pilbeam, Johanson, and Leakey, controversy and debate have always surrounded the search for our earliest ancestors. Most authorities now agree that many important questions concerning ape and human ances­try will be solved by further investigation of the hominoid primates of the Miocene and Pliocene, and through exacting comparative anatomical and biomolecular analyses of their living representatives. Indeed, these studies will yield a better understanding of (1) the cladogenesis (branching order) of the hominoid primates, (2) the morphotype (structural components) of the last common ancestor of humans and the living apes, (3) the timing and geographical placement of the hominid-pongid (human-ape) divergence and (4) the adaptive nature and probable scenario for the initial differentiation of hominids from pongids.

In this context we organized a symposium entitled "Miocene Hominoids and New Interpretations of Ape and Human Ancestry" which met July 2-4, 1980, in Florence, Italy,just prior to the convening of the VIII Congress ofthe International Primatological Society. This Pre-Congress Symposium was at­tended by a small group of anthropologists, anatomists, biochemists, ecolo­gists, and paleontologists, all dedicated to a reanalysis of the evolutionary position and a more parsimonious interpretation of the Miocene hominoids vis a vis modern apes and humans. The symposium was held in the 18th­Century Tribuna di Galileo of the Istituto di Zoologia. Under the arched marble columns of this magnificent chamber, symposium participants were able to address specifically some or all of the points mentioned above. From this symposium a consensus was reached concerning a rapprochement be­tween the formerly irreconcilable biomolecular view of human origins and the paleontological view. This rapprochement was based on a more liberal pre­sentation of the "molecular clock hypothesis" and on the view that no known Miocene hominoid including Ramapithecus could be considered a hominid, that is, an exclusively human ancestor.

xiii

xiv PREFACE

The Pre-Congress Symposium held in Florence was a rather small gathering, representing less than half of the contributors to this volume. Yet we feel that events at this symposium in many ways foreshadowed develop­ments and trends in the field of paleoanthropology still underway. With the symposium behind us we began in earnest to assemble contributions for the present volume. About this time a number of additional leading authorities, who had not been able to attend the symposium, agreed to contribute chap­ters to the volume. We also decided to enlarge the scope of the volume to include contributions on Oligocene hominoid ancestors and on Plio-Pleisto­cene hominoid and hominid descendants. The central theme of the volume, however, remained the Hominoidea of the Miocene.

The 30 chapters that follow approach the subject of ape and human ancestry from a wide variety of perspectives. Some of the topics covered include: biomolecular studies, craniofacial anatomy, cladistic methodologies, dental anatomy and histology, dental metrics, dietary adaptations, em­bryology, geochronology, karyology, paleoecology, paleomammalogy, paleo­neurology, parasitology, postcranial anatomy and locomotor adaptations, postcranial morphometrics, Popperian historicoscientific analyses, traditional descriptive analyses, and zoogeography. Whether or not our understanding of ape and human ancestry is further elucidated by this broad treatment is left for the reader to decide. Certainly though, the sheer quantity of data present­ed in this volume should be of use to researchers for many years to come.

Chapters 1 and 2 provide an introductory background to the remaining chapters of the volume. As the authors of Chapter 1, we attempt to present an even-handed, slightly historical overview of hominoid phyletics without trying to anticipate the many new perspectives proposed by contributors to this volume. Chapter 2, on the other hand, provides an in-depth review and syn­thesis of the geochronology and zoogeographic relationships of Miocene hominoids, which serves as a useful backdrop to many of the succeeding chapters. Finally in Chapter 30, RCL presents a summary viewpoint of ape and human ancestry based primarily on the contributions to this volume. In the time since this volume went to press new specimens have been recognized from several Eurasian and East African localities, which may already force revision of some conclusions presented herein. Clearly, at the present time, the field of hominoid evolution is so rapidly changing that no volume can stand as a complete summary for very long. Our hope is that each new volume and each new major revision will take the field one step closer to a new paradigm of ape and human ancestry that will better resist attempts to fal­sify it.

R. L. Ciochon R. S. Corruccini

Acknowledgments

Most cooperative scientific endeavors such as an edited volume owe their beginning in concept to a single idea or suggestion of one individual. In this regard, we express our sincere thanks to W. P. Luckett, who suggested the concept for this volume to RLC one cold December evening in 1978, while we waited at the Torino Airport. That suggestion grew into a 30-chapter volume with 37 contributors. In the process a number of individuals provided invalu­able scientific stimuli. We acknowledge R. L. Bernor, E. Delson, J. G. Fleagle, R. F. Kay, and T. D. White for this role. Eric Delson deserves a special note of thanks for suggesting several contributions to this volume, which otherwise would not have been included, and for putting up with innumerable queries concerning bibliographic and other details. Reviewing our graduate years, we would like to thank our major professors, F. Clark Howell and S. L. Wash­burn, for introducing us in the early 1970s to the field of Miocene hominoid evolution, therefore providing the conceptual framework for this volume. We also wish to thank the following students: Paula Atchison, Al Foster, Enita Mullens, Joy Myers, and Donna Ryan, for invaluable technical and editorial assistance. Ms. Myers, now in the graduate program at Kent State University, deserves special mention for having assisted at every stage during the prepara­tion of this volume and for single-handedly compiling all of the indexes. J. L. Simpson, G. M. Addington, and K. W. Culpepper of the Cartographic LabOIa­tory, 'Department of Geography and Earth Sciences, at the University of North Carolina at Charlotte are gratefully acknowledged for preparation of much of the artwork and photography in this volume. We also acknowledge the assistance of other staff members at UNCC, such as G. Ballard, D. Carter, and A. Salvo, and similarly thank T. Thomas from the Department of Anthropol­ogy at Southern Illinois University. For providing photographs used in the jacket illustration we thank P. J. Andrews and the British Museum of Natural History, R. F. Kay and the Duke Primate Center, W. H. Kimbel and the Cleveland Museum of Natural History, D. R. Pilbeam, W. Sacco and Harvard University, and A. C. Walker and the National Museum of Kenya. The plan­ning and production of the volume could not have taken place without the continued support and assistance of Kirk Jensen, Victoria Craven, and

xv

xvi ACKNOWLEDGMENTS

juiliana Newell Ayoub at Plenum. On behalf of all the contributors we whole­heartedly thank them. With regard to the Pre-Congress Symposium held in Florence, Italy, we thank A. B. Chiarelli, Director of the VIII Congress of the International Primatological Society, for his organizational efforts, and B. Lan­za, Director of the Istituto di Zoologia in Florence, for making the Tribuna di Galileo available for our symposiUJ;n. Financial support for symposium par­ticipants was provided by an L. S. B. Leakey Foundation grant to RLC, a NATO Fellowship to RSC, and additional funds from the National Science Foundation Iriternational Travel Support Program and the UNCC Interna­tional Studies Program. We also gratefully acknowledge the L. S. B. Leakey Foundation for granting additional research funds utilized throughout this volume's preparation. Finally we thank Francine Berkowitz of the Smithso­nian Foreign Currency Program for providing both moral and financial sup­port which greatly contributed to the completion of this work.

RLC RSC

Contents

I. PALEONTOLOGICAL AND GEOLOGICAL BACKGROUND

1 Overview of Ape and Human Ancestry: Phyletic Relationships of Miocene and Later

Hominoidea 3 R. S. Corruccini and R. L. Ciochon

2 Geochronology and Zoogeographic Relationships of Miocene Hominoidea .21

R. L. Bernor

II. EVIDENCE FROM MOLECULAR BIOLOGY AND COMPARATIVE ANATOMY

3 The Bearing of Molecular Data on the Cladogenesis and Times of Divergence of

Hominoid Lineages 67 M. Goodman, M. L. Baba, and L. L. Darga

4 A Model of Chromosome Evolution and Its Bearing on Cladogenesis in the

Hominoidea 87 L. L. Mai

5 Apes, Humans, and Molecular Clocks: A Reappraisal 115

J. E. Cronin

Appendix: Retrospective on Hominoid Macromolecular Systematics 137 V. M. Sarich

xvii

xviii CONTENTS

6 Cladistics and the Classification of the Great Apes 151

A. G. Kluge

III. EVIDENCE FROM CRANIODENTAL MORPHOLOGY

7 New Interpretations of the Phyletic Position of Oligocene Hominoids 181

J. G. Fleagle and R. F. Kay

8 Maxillofacial Morphology of Miocene Hominoids from Africa and Indo-

Pakistan 211 S. C. Ward and D. R. Pilbeam

9 A Reconsideration of the Endocast of Proconsul africanus: Implications for Primate

Brain Evolution 239 D. Falk

10 The Enamel of Neogene Hominoids: Structural and Phyletic Implications 249

D. G. Gantt

IV. EVIDENCE FROM POSTCRANIAL MORPHOLOGY

11 Locomotor Adaptations of Oligocene and Miocene Hominoids and Their Phyletic

Implications 301 J. G. Fleagle

12 New Postcranial Fossils of Proconsul africanus and Proconsul nyanzae 325

A. C. Walker and M. Pickford

13 The Wrist of Proconsul africanus and the Origin of Hominoid Postcranial

Adaptations 353 H. M. McHenry and R. S. Corruccini

CONTENTS xix

14 Miocene Hominoid Discoveries from Rudabanya: Implications from the Postcranial

Skeleton 369 M. E. Morbeck

15 Miocene Hominoid Postcranial Morphology: Monkey-like, Ape-like, Neither, or

Both? 405 M. D. Rose

V. EVIDENCE FROM PALEOENVIRONMENTAL STUDIES

16 Sequence and Environments of the Lower and Middle Miocene Hominoids of

Western Kenya 421 M. Pickford

17 The Natural History of Sivapithecus 441

P. J. Andrews

18 Facts and Fallacies Concerning Miocene Ape Habitats 465

A. Kortlandt

VI. DESCRIPTIVE ANALYSES OF SIWALIK MIOCENE HOMINOIDS

19 The Significance of Hitherto Undescribed Miocene Hominoids from the Siwaliks of

Pakistan in the Senckenberg Museum, Frankfurt 517 G. H. R. von Koenigswald

20 Miocene Hominoid Primate Dental Remains from the Siwaliks of

Pakistan 527 R. Dehm

XX CONTENTS

21 Significance of Recent Hominoid Discoveries from the Siwalik Hills of

India 539 S. R. K. Chopra

22 Historical Notes on the Geology, Dating and Systematics of the Miocene Hominoids

of India 559 K. N. Prasad

VII. ASSESSMENTS OF THE MIO-PLIOCENE EVIDENCE

23 A Reassessment of the Relationship between Later Miocene and Subsequent

Hominoidea 577 R. F. Kay and E. L. Simons

24 Phyletic Relationships of Miocene Hominoids and Higher Primate

Classification 625 L. de Bonis

25 Ramapithecus and Human Origins: An Anthropologist's Perspective of Changing

Interpretations 651 M. H. Wolpoff

26 Ramapithecus and Pan paniscus: Significance for Human Origins 677

A. L. Zihlman and J. M. Lowenstein

27 Toward the Resolution of Discrepancies between Phenetic and Paleontological Data

Bearing on the Question of Human Origins 695 L. O. Greenfield

CONTENTS xxi

28 Morphological Trends and Phylogenetic Relationships from Middle Miocene

Hominoids to Late Pliocene Hominids 705 N. T. Boaz

29 Australopithecus africanus: Its Phyletic Position Reconsidered 721

T. D. White, D. C. Johanson, and W. H. Kimbel

VIII. CONCLUDING REMARKS AND SUMMARY COMMENTS

30 Hominoid Cladistics and the Ancestry of Modern Apes and Humans: A Summary

Statement 781 R. L. Ciochon

Author Index 845

Taxonomic Index 861

Specimen Index 873

Subject Index 879

Guide to Specimen Abbreviations

The following is a list of alphabetic abbreviations of the institutional and/or locality designations of fossil specimens discussed in the chapters of this vol­ume. Since no key to specimen abbreviations is systematically presented in each chapter, this master list has been drawn up to aid readers. These same abbreviations are used in the index to specimens which appears at the end of this volume.

A.L.

AMNH

BMNHM

BP BSPhG

CGM

CYP

DPC

GSI GSI D­GSP KA

KNM­BC BN CA

Afar Locality, Hadar, Afar Triangle, Northeastern Ethiopia

American Museum of Natural History, New York, New York, U.S.A.

British Museum (Natural History), Mammal Division, London, United Kingdom

Bursa, Pa§alar Locality, Northwest Anatolia, Turkey Bayerische Staatssammlung fiir PaHiontologie und

historische Geologie, Munich, West Germany Cairo Geological Museum, Geological Surgey of Egypt,

Cairo, Egypt Chandigarh-Y ale Project, Panjab, University,

Chandigarh, India Duke Primate Center, Duke University, Durham, North

Carolina, U.S.A. Geological Survey of India, Calcutta, India Geological Survey of India, D Series, Calcutta, India Geological Survey of Pakistan, Quetta, Pakistan Kromdraai, Northern Transvaal, Republic of Soud;.

Africa Kenya National Museum, Nairobi, Kenya

Baringo, Chemeron Baringo, Ngorora Chamtwara Member, Koru

xxiii

xxiv SPECIMEN ABBREVIATIONS

CH ER IT KA KO KP LT MB ME MJ MW RU SO X

L.H.­MCZ

MLD

MTA

O.H. Omo

ONGC PA

PUA

RPI

Rud SK

SNM

STS

STW

TM UMP USNM

YPM

Chesowanja East Rudolf Fort Ternan Karungu Koru Kanapoi Lothagam Maboko Island Meswa Bridge M<tii",a Mfwangano Island Rusinga Island Songhor Unknown

Laetoli Hominid -, Laetolil Beds, Northern Tanzania Museum of Comparative Zoology, Harvard University,

Cambridge, Massachusetts, U.S.A. Makapansgat Limeworks Dump, Northern Transvaal,

Republic of South Africa Maden Tetkik ve Arama Enstitiisu (Mineral Research

and Exploration Institute), Ankara, Turkey Olduvai Hominid, Olduvai Gorge, Northern Tanzania Omo Locality, Shungura Formation, Lower Omo Basin,

Southern Ethiopia Oil and Natural Gas Commission, Dehra Dun, India Paleoanthropology Collection, Institute of Vertebrate

Paleontology and Paleoanthropology, Academia Sinica, Beijing, China

Panjab University, Anthropology Department, Chandigarh, India

Ravin de la Pluie (Rain Ravine), Lower Macedonia, Greece

Rudabanya, Northeastern Hungary Swartkrans, Northern Transvaal, Republic of South

Africa Staatliches Museum fur Naturkunde, Stuttgart, West

Germany Sterkfontein (Type Site) Northern Transvaal, Republic

of South Africa Sterkfontein (West Pit) Northern Transvaal, Republic of

South Africa Transvaal Museum, Pretoria, Republic of South Africa Uganda Museum, Primate Collection, Kampala, Uganda United States National Museum, Smithsonian Institution,

Washington, D.C., U.S.A. Yale Peabody Museum, Yale University, New Haven,

Connecticut. U.S.A.