Table Of ContentAnthropoid Origins
ADVA NCES IN PRIMAT OLOGY
Series Editors:
lOHN G. FLEAGLE
State University of New York at Stony Brook
Stony Brook, New York
R. D. E. MACPHEE
Ameriean Museum of Natural History
New York, New York
Editorial Board:
lOHN M. ALLMAN, California Institute of Technology, Pasadena, California
RUSSELL L. CIOCHON, University of lowa, lowa City, lewa
FRAN<;:OISEJOUFFROY, Museum National d'Histoire Naturelle, Paris, France
W. PATRICK LUCKETT, University of Puerto Rieo, Sanluan, Puerto Rieo
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
CONTINENTA L 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
NURSERY CARE OF NONHUMAN PRIMATES
Edited by Gerald C. Ruppenthal
PRIMATES AND THEIR RELATIVES IN PHYLOGENETIC
PERSPECTIVE
Edited by Ross D. E. MacPhee
SENSORY SYSTEMS OF PRIMATES
Edited by Charles R. Noback
SIZE AND SCALING IN PRIMATE BIOLOGY
Edited by William L. Jungers
SPECIES, SPECIES CONCEPTS, AND PRIMATE EVOLUTION
Edited by William H. Kimbel and Lawrence B. Martin
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.
Anthropoid Origins
Edited by
JOHN G. FLEAGLE
State University of New York at Stony Brook
Stony Brook, New York
and
RICHARD F. KAY
Duke University Medical Center
Durham, North Carolina
Springer Science+Business Media, LLC
Llbrary of Congress Cataloglng-ln-Publicatlon Data
Anthropoid origins I edited by John G. Fleagle and Riehard F. Kay.
p. e •. -- (Advances in primatologyl
Revised papers froN a conferenee held at Duke University. May
1992.
Ineludes bibliographieal referenees and index.
1. Primates--Evolution--Congresses. 2. Primates. Fossil-
-Congresses. I. Fleagle. John G. 11. Kay. Riehard F.
Irr. Series.
OL737.P9A64 1994
599.80438--dc20 94-43006
CIP
ISBN 978-1-4757-9199-0 ISBN 978-1-4757-9197-6 (eBook)
DOIlO.1007/978-1-4757-9197-6
©1994 Springer Science+Business Media New York
Originally published by Plenum Press, New York in 1994.
Softcover reprint ofthe hardcover 1s t edition 1994
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 Elwyn L. Simons
Who over the past thirty-five years
invented paleoprimatology as a
discipline, and whose ongoing
discoveries in Egypt continue
to set the agenda for studies of
anthropoid origins
Contributors
K. Christopher Beard John G. Fleagle
Section of Vertebrate Paleontology Department of Anatomical Sciences
Carnegie Museum of Natural History School of Medicine
Pittsburgh, Pennsylvania 15213 Health Sciences Center
State University of New York
Thornas M. Bown Stony Brook, New York 11794
Paleontology and Stratigraphy Branch
U.S. Geological Survey Susan M. Ford
Denver, Colorado 80225 Department of Anthropology
Southern Illinois University
Malt Cartrnill Carbondale, Illinois 62901
Department of Biological
Anthropology and Anatomy Jens Lorenz Franzen
Duke University Forschungsinstitut Senckenberg
Durham, North Carolina 27710 D-60325 Frankfurt am Main, Germany
Prithijit S. Chatrath Daniel L. Gebo
Duke University Primate Center Department of Anthropology
Durham, North Carolina 27706 Northern Illinois University
DeKalb, Illinois 60115
Russell L. Ciochon
Departments of Anthropology and Philip D. Gingerieh
Pediatric Dentistry Museum of Paleontology
University of Iowa University of Michigan
Iowa City, Iowa 52242 Ann Arbor, Michigan 48109
Herbert H. Covert Mare Godinot
Department of Anthropology Laboratoire de Paleontologie
University of Colorado Institut des Sciences de l'Evolution
Boulder, Colorado 80309-0233 Universite Montpellier II
F-34095 Montpellier Cedex 5, France
Marian Dagm"to
Department of Cell and Molecular Patrieia A. Holroyd
Biology Department of Biological
Northwestern University Medical Anthropology and Anatomy
School Duke University
Chicago, Illinois 60611, and Durham, North Carolina 27710, and
Department of Mammalogy U.S. Geological Survey
American Museum of Natural History Denver, Colorado 80225
New York, New York 10024
vii
viii CONTRIBUTORS
William L. Hylander Kenneth D. Rose
Department of Biological Department of Cell Biology and
Anthropology and Anatomy Anatomy
Duke University Medical Center Johns Hopkins University School of
Durham, North Carolina 27710 Medicine
Baltimore, Maryland 21205
Richard F. Kay
Department of Biological Callum Ross
Anthropology and Anatomy Department of Biological
Duke University Anthropology and Anatomy
Durham, North Carolina 27710 Duke Universitv
Durham, Nortl~ Carolina 27705
Mary C. Maas Present address:
Department of Biological Department of Anatomical Sciences
Anthropology and Anatomy School of Medicine
Duke University Health Sciences Center
Durham, North Carolina 27710 State University of New York
Stony Brook, New York 11794
R. D. E. MacPhee
Department of Mammalogy Elwyn L. Simons
American Museum of Natural History Duke Primate Center
New York, New York 10024 Duke University
Durham, North Carolina 27705
D. Tab Rasmwsen
Department of Anthropology Blythe A. Williams
Washington University Department of Anthropology
St. Louis, Missouri 63130 University of Colorado
Boulder, Colorado 80309
Matthew J. Ravosa Present address:
Department of Biological Department of Biological
Anthropology and Anatomy Anthropology and Anatomy
Duke University Medical Center Duke University
Durham, North Carolina 27710 Durham, North Carolina 27710
Present address:
Department of Ceil and Molecular
Biology
Northwestern University Medical
School
Chicago, IIIinois 60611-3008, and
Department of Zoology
Division of Mammals
Field Museum of Natural History
Chicago, IIIinois 60605-2496
Preface
This volume brings together information about recent discoveries and current
theories concerning the origin and early evolution of anthropoid primates
monkeys, apes, and humans. Although Anthropoidea is one of the most dis
tinctive groups of living primates, and the origin of the group is a frequent
topic of discussion in the anthropological and paleontological literature, the
topic of anthropoid origins has rarely been the foeus of direct discussion in
primate evolution. Rather, diseussion of anthropoid origins appears as a ma
jor side issue in volumes dealing with the origin of platyrrhines (Ciochon and
Chiarelli, 1980), in discussions about the phylogenetic position of Tarsius, in
descriptions of early anthropoid fossils, and in descriptions and revisions of
various fossil prosimians. As a result, the literature on anthropoid origins has
a long history of argument by advocacy, in which scholars with different views
have expounded individual theories based on a small bit of evidence at hand,
often with little consideration of alternative views and other types of evidence
that have been used in their support. This type of scholarship struck us as a
relatively unproductive approach to a critical issue in primate evolution. We
hoped that by bringing together scholars with expertise in different areas of
primate evolution and often divergent views of this issue, we could generate a
common focus and broader awareness, in order to sort out areas of general
agreement, areas of great disagreement, and areas of acknowledged igno
rance. In doing so we would be in a better position to either resolve the
quest ion of anthropoid origins or at least set the stage for more fruitful
discussion of the topic in future years.
To this end, in May 1992, we joined with Professor Elwyn Simons, Scien
tific Director ofthe Duke University Primate Center, to organize a conference
and workshop on Anthropoid Origins at Duke University. Through four days
of presenting papers, diseussing anthropoid origins, examining new fossils,
and watching lemurs, all ofthe participants gained a better knowledge ofboth
the available fossil material, much of it unpublished at the time, and the views
of the other participants. Following the conference, many of the participants
ix
X PREFACE
wrote up and submitted manuscripts on various aspects of anthropoid ori
gins; these were sent to other participating scientists as weIl as "outside"
scholars for peer revieW. The manuscripts were revised, often drasticaIly,
before they were accepted for publication in the volume. As a result the
papers contained here reflect the combined input of many researchers and
demonstrate real progress in addressing the question of anthropoid origins.
The problem of anthropoid origins is not a single issue, but a wide range
of issues. These obviously include the phylogenetic origin of anthropoids
(What is the sister taxon of anthropoids?), the geographical origin of the
group (Where did anthropoids first evolve?), and the less frequently ad
dressed problem of the adaptations of the earliest anthropoids (Did the origin
of anthropoids involve a major adaptive shift in primate evolution, or so me
"keystone adaptation"?). But, in addition, the question includes sorting out
the phylogenetic relationships (and adaptations) of an increasing diversity of
extinct species that appear to be near the border between prosimians and
anthropoids. While the origin of anthropoids may appear to be a single point
in a cladogram of living primates, the stern between the earliest anthropoid
and the subsequent divergence of modern anthropoid groups could, and
almost certainly did, contain dozens of extinct taxa possessing a variety of
combinations of "anthropoid" features. The anthropoid tree is much bushier
at the base than the diversity of modern taxa would indicate.
This volume addresses these many aspects of anthropoid origins from a
variety of perspectives. So me chapters consider anthropoid origins from the
perspective of the prosimians that preceded and perhaps gave rise to higher
primates. Some take an anthropoid perspective to describe the early anthro
poid radiation and "look back" to see where the group may have come from.
They examine hoth prosimians and anthropoids using cranial, dental, or
postcranial features. The authors have been encouraged to compare and
contrast their results with those of others in the volume so that both individu
ally and collectively they strive for some degree of synthesis.
This volume has benef'ited from the contributions of many persons and
institutions. Funding for the conference was provided in part by the LSB
Leakey Foundation, The Wenner-Gren Foundation, The National Science
Foundation, Duke University, and the State University of New York. The
conference and the volume would not have been possible without the many
contributions of Dr. Elwyn Simons, who as scientist, host, and mentor to many
of the participants laid the base for most of the research included in the
volume. Ken Glander, Prithijit Chatrath, Friderun Ankel-Simons, Todd Rae,
Mario Gagnon, Irene Lofstrum, Gay Gaster, and Rachael Hougam were in
strumental in making the conference a success. Todd Rae, Walter Hartwig,
Joan KeIly, and Kaye Reed made herculean efforts in putting together the
chapters that make up the volume. Many of our colleagues aided in reviewing
the manuscripts. Most of all, however, this volume reflects the willingness of
our colleagues to share their time, their thoughts, their diverse skills, and
