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RECONSTRUCTING BEHAVIOR IN THE PRIMATE FOSSIL 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 University of New York at Stony Brook
Stony Brook, New York

and

Carel P. van Schaik
Department of Biological Anthropology and Anatomy
Duke University Medical Center
Durham, North Carolina

Kluwer 2002

FROM THE BACK COVER

This volume brings together a series of papers that address the topic of reconstructing 
behavior in the primate fossil record.

Offered here are reviews of broad issues related to reconstructing various aspects of 
behavior-from diet to social systems-from different types of evidence including 
comparative analysis and models of adaptation, ontogenetic evidence, paleo-
environmental and paleo-community analysis, experimental functional analysis, and 
comparative socioecology. Authors review not only the types of evidence that can be 
used to reconstruct behavior, but also the limitations of the evidence, pointing to 
directions for future research.

Hopefully, 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 new approaches to the topic.

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. Hylander 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

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

9. The Adaptations of Branisella boliviana,
the Earliest South American Monkey             339
Richard F. Kay, Blythe A. Williams,
and Federico Anaya
   Introduction     339
   Branisella boliviana, the Earliest-Known Platyrrhine Primate     345
   Summary and Conclusions    362
   Appendix     364
   References     366

10. Ecomorphology and Behavior of
Giant Extinct Lemurs from Madagascar     371
William L. Jungers, Laurie R. Godfrey,
Elwyn L. Simons, Roshna E. Wunderlich,
Brian G. Richmond, and Prithijit S. Chatrath
   Introduction     371
   Body Size and Sexual Dimorphism     374
   Activity Cycles          376
   Oral Behaviors     381
   Positional Behavior     390
   Summary    401
   References    404

11. Conclusions: Reconstructing Behavior in the Fossil Record   413
J. Michael Plavcan, Richard F. Kay,
William L. Jungers, and Carel P. van Schaik
   Introduction      413
   The Comparative Approach      414
   Phylogeny      417
   Standard Error from Comparative Analyses      418
   Body Mass and Allometry      419
   Incomplete Extant Models       421
   Problems with Defining Behavior      422
   Multiple Lines of Evidence      424
   Conclusions      425
   References     426

Subject Index     429

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 overwhelming 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 increasingly rigorous.

We offer here a series of papers that review broad issues related to reconstructing various 
aspects of behavior from very different types of evidence. 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 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, paleoenvironmental and paleocommunity analysis, 
experimental functional analysis, and comparative 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 
reconstructions for extinct femurs 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 formulating 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

CONTRIBUTORS

Federico Anaya
Museo Nacional de Historia Naturale
La Paz, Bolivia

Prithijit S. Chatrath
Duke University Primate Center
Durham, North Carolina 27705

John G. Fleagle
Department of Anatomical Sciences
Health Sciences Center
State University of New York at Stony Brook
Stony Brook, New York 11794-8081

Laurie R. Godfrey
Department of Anthropology
University of Massachusetts
Amherst, Massachusetts 01003-9278

William L. Hylander
Department of Biological Anthropology and Anatomy
Duke University Medical Center
Durham, North Carolina 27710

Kirk R. Johnson
Department of Biological Anthropology and Anatomy
Duke University Medical Center
Durham, North Carolina 27710

William L. Jungers
Department of Anatomical Sciences
Health Sciences Center
State University of New York at Stony Brook
Stony Brook, New York 11794-8081

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

Charles A. Lockwood
Institute of Human Origins
Arizona State University
Tempe, Arizona 85287-4101

Charles L. Nunn
Department of Biological Anthropology and Anatomy
Duke University
Durham, North Carolina 27708-0383
Present address:
Section of Evolution and Ecology
University of California
Davis, California 95616

Andrew J. Petto
Division of Liberal Arts
University of the Arts
Philadelphia, Pennsylvania 19102-4994

WHERE TO ORDER

Kluwer Academic Publishers
Order Department
P.O. Box 358, Accord Station
Hingham, MA 02018-0358, USA

Tel: (781) 871-6600
Fax: (781) 871-6528
E-mail: kluwer@wkap.com
http://www.wkap.com

Price: $125.00  ISBN: 0-306-46604-X (hardbound)


Book received: 6-17-02
Posted date: 9-12-02