Table Of ContentReconstructing Behavior
in the Primate
Fossil Record
ADVANCES IN PRIMATOLOGY
Series Editors:
JOHN G. FLEAGLE
State University of New York at Stony Brook
Stony Brook, New York
R. D. E. MAcPHEE
American Museum of Natural History
New York, New York
Editorial Board:
JOHN M. ALLMAN, California Institute of Technology, Pasadena, California
RUSSEL L. CIOCHON, University of Iowa, Iowa City, Iowa
FRAN<;;OISE JOUFFROY, Museum National d'Histoire Naturelle, Paris, France
W. PATRICK LUCKETT, University of Puerto Rico, San Juan, Puerto Rico
LAWRENCE B. MARTIN, State University of New York at Stony Brook, Stony Brook, New York
CAREL P. VAN SCHAlK, Duke University, Durham, North Carolina
Current Volumes in the Series:
ANTHROPOID ORIGINS
Edited by John G. Fleagle and Richard F. Kay
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
FUNCTION, PHYLOGENY, AND FOSSILS: Miocene Hominoid Evolution and
Adaptations
Edited by David R. Begun, Carol V. Ward, and Michael D. Rose
NEW INTERPRETATIONS OF APE AND HUMAN ANCESTRY
Edited by Russell L. Ciochon and Robert S. Corroccini
NURSERY CARE OF NONHUMAN PRIMATES
Edited by Gerald C. Ruppenthal
PRIMATES AND THEIR RELATIVES IN PHYLOGENETIC PERSPECTIVE
Edited by Ross D. E. MacPhee
RECONSTRUCTING BEHAVIOR IN THE PRIMATE FOSSIL RECORD
Edited by J. Michael Plavcan, Richard F. Kay, William L. Jungers,
and Carel P. van Schaik
SIZE AND SCALING IN PRIMATE BIOLOGY
Edited by William L. Jungers
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.
Reconstructing Behavior
in the Primate
Fo ssil Record
Edited by
J.
MICHAEL PLAVCAN
Department of Anthropology
University of Arkansas
Fayetteville, Arkansas
RICHARD F. KAY
Department of Biological Anthropology and Anatomy
Duke University Medical Center
Durham, North Carolina
WILLIAM L. JUNGERS
Department of Anatomical Sciences
State Unviersity of New York at Stony Brook
Stony Brook, New York
and
CAREL P. SCHAIK
VAN
Department of Biological Anthropology and Anatomy
Duke University Medical Center
Durham, North Carolina
Springer Science+Business Media, LLC
Library of Congress Cataloging-in-PublicaLion Data
ReeonstrucLing behavior in the primate fossil record/edited by j. Miehael PlaH:an ... [et al.].
p. em. - (Advanees in primatology)
Includes bibliographical referenccs and index.
ISBN 978-1-4613-5507-6 ISBN 978-1-4615-1343-8 (eBook)
DOI 10.1007/978-1-4615-1343-8
1. Primates. Fossil. 2. Primates-Behavior. 3. Evolutionary paleobiology. 1. Plavt:an,j.
Michael, 1962- II. Advanees in primalOlogy (Kluwer Academic/ Plenum Publishers)
QE882.P7 R43 2002
569' .8-dc21
2001032668
ISBN 978-146\3-5507-6
(:2002 Springer Science+Business Media New York
OriginaJly published by Kluwer Academic! Plenum Publishers. New York in 2002
Softoo\"er rc:print oflhe hardcover ISI edition 2002
hup:I/_"W.wk.ap.nl/
10987654321
A C.I.P. record for this boole. is available from the Library of Congress
Ali rights resen·ed
No part of this boole. may be reproduced, slon:d in a retrieval s)"Stem, or transmitted in any form
or by any means, electronic, mechanical, phOlocopying, mieroflhning, recording, or olherwise,
without written permission from the Publisher
Contributors
Federico Anaya William L. Jungers
Museo Nacional de Historia Naturale Department of Anatomical Sciences
La Paz, Bolivia Health Sciences Center
State University of New York
Prithijit S. Chatrath at Stony Brook
Duke University Primate Center Stony Brook, N ew York 11794-8081
Durham, North Carolina 27705
Richard F. Kay
Department of Biological Anthropology
John G. Fleagle
and Anatomy
Department of Anatomical Sciences
Duke University Medical Center
Health Sciences Center
Durham, North Carolina 27710
State University of New York
at Stony Brook
Charles A. Lockwood
Stony Brook, New York 11794-8081
Institute of Human Origins
Arizona State University
Laurie R. Godfrey
Tempe, Arizona 85287-4101
Department of Anthropology
University of Massachusetts
Charles L. Nunn
Amherst, Massachusetts 01003-9278
Department of Biological Anthropology
and Anatomy
William L. Rylander Duke University
Department of Biological Anthropology Durham, North Carolina 27708-0383
and Anatomy Present address:
Duke University Medical Center Section of Evolution and Ecology
Durham, North Carolina 27710 University of California
Davis, California 95616
Kirk R. Johnson
Department of Biological Anthropology Andrew J. Petto
and Anatomy Division of Liberal Arts
Duke University Medical Center University of the Arts
Durham, North Carolina 27710 Philadelphia, Pennsylvania 19102-4994
v
vi CONTRIBUTORS
J. Michael Plavcan Callum F. Ross
Department of Anatomy Department of Anatomical Sciences
New York College of Osteopathic Health Sciences Center
Medicine State University of New York
Old Westbury, New York 11568 at Stony Brook
Present address: Stony Brook, New York 11794-8081
Department of Anthropology Elwyn L. Simons
University of Arkansas Duke University Primate Center
Fayetteville, Arkansas 72701 Durham, North Carolina 27705
Michael R. Sutherland
Matthew J. Ravosa Statistical Consulting Center
Department of Cell and Molecular University of Massachusetts, Amherst
Biology Amherst, Massachusetts 01003-9337
Northwestern University Medical School
Chicago, Illinois 60611-3008
Peter Ungar
and Department of Anthropology
Department of Zoology
University of Arkansas
Division of Mammals Fayetteville, Arkansas 72701
Field Museum of Natural History
Chicago, Illinois Carel P. van Schaik
Department of Biological Anthropology
and Anatomy
Kaye E. Reed
Duke University
Institute of Human Origins and
Durham, North Carolina 27708-0383
Department of Anthropology
Arizona State University
Christopher J. Vinyard
Tempe, Arizona 85281-4101
Department of Biological Anthropology
and
and Anatomy
Bernard Price Institute of
Duke University Medical Center
Palaeontological Research
Durham, North Carolina 27710
University of the Witwatersrand
Johannesburg, South Africa
Blythe A. Williams
Department of Biological Anthropology
Brian G. Richmond and Anatomy
Department of Anthropology Duke University Medical Center
George Washington University Durham, North Carolina 27710
Washington DC 20052
Present address: Roshna E. Wunderlich
Department of Anthropology Department of Biology
University of Illinois James Madison University
Urbana, Illinois 61801 Harrisburg, Virginia 22807
Preface
This volume brings together a series of papers that address the topic of
reconstructing behavior in the primate fossil record. The literature devoted to
reconstructing behavior in extinct species is ovelWhelming and very diverse.
Sometimes, it seems as though behavioral reconstruction is done as an
afterthought in the discussion section of papers, relegated to the status of
informed speculation. But recent years have seen an explosion in studies of
adaptation, functional anatomy, comparative sociobiology, and development.
Powerful new comparative methods are now available on the internet. At the
same time, we face a rapidly growing fossil record that offers more and more
information on the morphology and paleoenvironments of extinct species.
Consequently, inferences of behavior in extinct species have become better
grounded in comparative studies of living species and are becoming increas
ingly rigorous.
We offer here a series of papers that review broad issues related to
reconstructing various aspects of behavior from very different types of evi
dence. We hope that in so doing, the reader will gain a perspective on the
various types of evidence that can be brought to bear on reconstructing
behavior, the strengths and weaknesses of different approaches, and, perhaps,
new approaches to the topic. We define behavior as broadly as we can
including life-history traits, locomotion, diet, and social behavior, giving the
authors considerable freedom in choosing what, exactly, they wish to explore.
Notably, we deliberately exclude issues related to later human evolution and
behavior, especially with regard to archeological evidence. There are several
extensive works from the archeological literature that deal well with the unique
problems and opportunities available to students of the human fossil record.
We asked authors to address topics with a broad approach. We especially
encouraged authors to review not only the types of evidence that can be used
to reconstruct behavior, but also the limitations of the evidence. We feel that
it is important to understand both what can, and cannot, be said about the
behavior of extinct species. Our intent is not to cast behavioral reconstruction
in a cynical light, but to emphasize the weaknesses of behavioral reconstruction
vii
viii
PREFACE
as a basis for further research. We hope that by understanding both the
strengths and weaknesses of behavioral reconstructions, future research will
create a much more rigorous framework for understanding the adaptations
and behaviors of extinct animals.
We present first a series of chapters that systematically address basic issues
and types of evidence that can be used to infer behavior. These include
comparative analysis and adaptation, ontogenetic evidence, paleoenvironmen
tal and paleocommunity analysis, experimental functional analysis, and com
parative socioecology. This is followed by two broad reviews of evidence for
diet and social systems in primates - two of the most commonly inferred
behaviors for extinct species. We end with reviews of behavioral reconstruc
tions for extinct lemurs and Branisella boliviana, each of which illustrates how
widely different types of evidence converge to paint a picture of the behavior
and adaptations of extinct species.
We hope that students and professionals find this book interesting and
useful for thinking about how to reconstruct behavior in the fossil record. We
thank the authors for their thoughtful contributions and especially their
patience in producing this volume. The difficulty of reviewing such a broad
topic in an accessible fashion offers a real challenge, which all of the authors
successfully met. This volume was initially inspired by a conference at Duke
University sponsored by the Leakey Foundation, whom we thank for their
generous support. John Fleagle and Ross MacPhee helped greatly in formula
ting the structure of the volume. Finally, we thank the many external reviewers
of the papers, who provided thoughtful and insightful comments on the
manuscripts, and greatly improved the quality of the volume.
J. Michael Plavcan
Richard F. Kay
William L. Jungers
Carel P. van Schaik
Old Westbury, New York
Durham, North Carolina
Stony Brook, New York
Durham, North Carolina
Contents
1
Adaptation and Behavior in the Primate Fossil Record 1
Callum F. Ross, Charles A. Lockwood, John G. Fleagle,
and William L. Jungers
Introduction 1 Defining Adaptation 2 IdentifYing Adaptations 4
How Can Fossil Taxa Be Used to Study Adaptation? 32 Summary and
Conclusions 34 References 35
2
Functional Morphology and In Vivo Bone Strain Patterns in the Craniofacial
Region of Primates: Beware of Biomechnical Stories about Fossil
Bones 43
William L. Rylander and Kirk R. Johnson
Introduction 43 In Vivo Bone Strain Patterns 47 Discussion 53
Conclusions 67 References 68
3
On the Interface between Ontogeny and Function 73
Matthew J. Ravosa and Christopher J. Vinyard
Introduction 73 Ontogeny as a Criterion of Subtraction 74
Biomechanical Scaling and Functional Equivalence 87 Heterochrony 97
Conclusions 104 References 105
ix
X CONTENTS
4
Dental Ontogeny and Life-History Strategies:
The Case of the Giant Extinct Indroids of Madagascar 113
Laurie R. Godfrey, Andrew J. Petto, and Michael R. Sutherland
Introduction 113 Development and the Reconstruction of the Behavior and Life
Histories of Extinct Primates: Approaches and Methodologies 114 Development and
the Reconstruction of Behavior in Extinct Primates: An Example 122
Conclusions 148 Appendix 150 References 153
5
A Comparative Approach to Reconstructing the Socioecology of Extinct
Primates 159
Charles L. Nunn and Carel P. van Schaik
Introduction 159 Methods 163 Results 166 Worked Examples 184
Discussion 190 Conclusions 198 Appendix 200 References 211
6
The Use of Paleocommunity and Taphonomic Studies in Reconstructing
Primate Behavior 217
Kaye E. Reed
Introduction 217 The Comparative Method and Actualistic Studies in
Paleoecology 220 Fossil Assemblages 222 Community Comparisons 234
Behavioral Ecology of Primates in Extant and Makapansgat Communities 242 Primate
Interactions and Behavior Based on Accumulating Agents 247 Integration of
Community Comparisons 249 Evolution, Taphonomy, and Community
Paleoecology 254 References 255
7
Reconstructing the Diets of Fossil Primates 261
Peter Ungar
Introduction 261 Adaptive Signals for Diet in Primates 262 Nonadaptive Signals
for Diet in Primates 276 Discussion 283 Summary and
Conclusions 286 References 288
8
Reconstructing Social Behavior from Dimorphism in the Fossil Record 297
J. Michael Plavcan
Introduction 297 Data 301 Behavioral Variables 302 Relations between
Dimorphism and Behavioral Estimates 306 Relations between Relative Canine Size
and Competition Classifications 317 Relations between Dimorphism and Other
Variables 321 Summary and Conclusions 332 References 333
