Table Of ContentComparative Biology and
Evolutionary Relationships
of Tree Shrews
ADVANCES IN PRIMATOLOGY
Series Editors:
W. PATRICK LUCKETT
Creighton University School of Medicine
Omaha, Nebraska
CHARLES R. NOBACK
Columbia University
New York, New York
Editorial Board:
JOHN F. EISENBERG F.A.JENKINS,Jr.
Smithsonian Institution Harvard University
Washington, D.C. Cambridge, Massachusetts
MORRIS GOODMAN FREDERICK S. SZALAY
Wayne State University School of Medicine Hunter College
Detroit, Michigan New York, New York
mE 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
A Continuation Order Plan is available for tbis 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.
Comparative Biology and
Evolutionary Relationships
of Tree Shrews
Edited by
w.
Patrick Luckett
Creighton University School of Meaiclne
Omaha, Nebraska
Plenum Press . New York and London
Library of Congress Cataloging in Publication Data
Main entry under title:
Comparative biology and evolutionary relationships of tree shrews.
(Advances in primatology)
Includes bibliographical references and index.
1. Tupaiidae-Evolution. 2. Mammals-Evolution. I. Luckett, Winter Patrick.
II. Series.
QL737.P968C65 599.3'3 80-19824
ISBN 978-1-4684-1053-2
ISBN 978-1-4684-1053-2 ISBN 978-1-4684-1051-8 (eBook)
001 10. 1007/978-1-4684-1051-8
© 1980 Plenum Press, New York
Softcover reprint of the hardcover 1s t edition 1980
A Division of Plenum Publishing Corporation
227 West 17th Street, New York, N.Y. 10011
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
Contributors
Percy M. Butler Gerrell Drawhorn
Department of Zoology Department of Anthropology
Royal Holloway College University of California, Davis
Englefield Green, Surrey Davis, California
England
Morris Goodman
Department of Anatomy
C. B. G. Campbell
Wayne State University School
Department of Medical
of Medicine
Neurosciences
Detroit, Michigan
Division of Neuropsychiatry
Walter Reed Army Institute of
Research Louis L. Jacobs
Washington, D. C. Department of Geology
Museum of Northern Arizona
Flagstaff, Arizona
Matt Cartmill
Department of Anatomy
W. Patrick Luckett
Duke University Medical Center
Department of\Anatomy
Durham, North Carolina
Creighton University
Omaha, Nebraska
John E. Cronin
Department of Anthropology R. D. E. MacPhee
Peabody Museum Department of Anatomy
Harvard University Duke University Medical Center
Cambridge, Massachusetts Durham, North Carolina
Howard Dene Genji Matsuda
Department of Anatomy Department of Biochemistry
Wayne State University School Nagasaki University School of
of Medicine Medicine
Detroit, Michigan Nagasaki, Japan
v
vi CONTRIBUTORS
J.
Michael Novacek Vincent M. Sarich
Department of Zoology Departments of Anthropology
San Diego State University and Biochemistry
San Diego, California University of California
Berkeley, California
William Prychodko Frederick S. Szalay
Department of Biology Department of Anthropology
Wayne State University Hunter College, CUNY
Detroit, Michigan New York, New York
Preface
Tree shrews are small-bodied, scansorial, squirrel-like mammals that occupy
a wide range of arboreal, semi-arboreal, and forest floor niches in Southeast
Asia and adjacent islands. Comparative aspects of tree shrew biology have
been the subject of extensive investigations during the past two decades. These
studies were initiated in part because of the widely accepted belief that tupaiids
are primitive primates, and, as such, might provide valuable insight into the
evolutionary origin of complex patterns of primate behavior, locomotion,
neurobiology, and reproduction. During the same period, there has been a
renewed interest in the methodology of phylogenetic reconstruction and in
the use of data from a variety of biological disciplines to test or formulate
hypotheses of evolutionary relationships. In particular, interest in the com
parative and systematic biology of mammals has focused on analysis of phy
logenetic relationships among Primates and a search for their closest relatives.
Assessment of the possible primate affinities of tree shrews has comprised an
important part of these studies, and a considerable amount of dental, cranio
skeletal, neuroanatomical, reproductive, developmental, and molecular evi
dence has been marshalled to either corroborate or refute hypotheses of a
special tupaiid-primate relationship. These contrasting viewpoints have re
sulted from differing interpretations of the basic data, as well as alternative
approaches to the evolutionary analysis of data.
The present volume was organized in order to evaluate the possible evo
lutionary relationships of tree shrews to primates and other eutherian mam
mals. Such analysis is deemed an essential prerequisite for the use of tree
shrews as possible models for the study of the evolutionary origin of various
primate organ systems. Assessment of the mammalian affinities of tree shrews
is limited because of the sparseness of the tupaiid fossil record (restricted at
present to the recently discovered Miocene specimens described by Jacobs in
this volume). In order to evaluate the possible mammalian affinities oftupaiids,
it is essential to attempt reconstruction of the morphotypic condition for each
organ system analyzed in Tupaiidae, Primates, Dermoptera, Chiroptera, Li
potyphla, Macroscelididae, and other eutherian taxa, because such analyses
vii
viii PREFACE
facilitate the recognition of convergent evolution of individual traits in these
systems. Reconstruction of the eutherian morphotype for selected features
was also undertaken by many contributors, and these hypothetical reconstruc
tions serve as focal points for identifying areas of disagreement in phylogenetic
analyses.
Contributors to the volume were requested to consider two interrelated
questions during their assessments of the possible evolutionary relationships
of tree shrews. (1) Are there uniquely derived biological attributes shared solely
by tupaiids and primates? (2) If tupaiids are not cladistically primates, are they
more closely related phyletically to primates, dermopterans, and chiropterans
(in a superordinal taxon Archonta) than they are to any other eutherians?
Following evaluations of the available dental, cranial, postcranial, neuroana
tomical, reproductive, developmental, and molecular evidence, the contribu
tors were in general agreement that tupaiids do not appear to share any
uniquely derived features with living or fossil primates that would warrant
inclusion of tree shrews in a monophyletic order Primates. Available biological
data suggest, instead, that tree shrews have evolved independently since at
least the early Tertiary, despite the fact that corroborating fossil evidence is
lacking. As proposed previously by Butler, recognition of the separate ordinal
status of tree shrews (as the order Scandentia) appears to be the best manner
of expressing such a long and independent evolutionary history.
There was less agreement among contributors concerning the possible
superordinal relationships of Scandentia. The available molecular and post
cranial evidence provides some support for a modified archontan hypothesis
of phylogenetic affinities among Scandentia, Primates, and Dermoptera, but
data evaluated from other organ systems provide little, if any, corroboration
for this postulate. As emphasized by Szalay and Drawhorn in this volume,
however, the available biological evidence does not corroborate any alternative
hypotheses of superordinal affinities of Scandentia. Future investigations of
tupaiid biology should provide additional data for testing the archontan
hypothesis.
Even if tree shrews are not considered to be members of the order Pri
mates, these fascinating mammals will remain the subject of extensive study
by primate biologists. Analyses of the evolutionary relationships of tree shrews
during the past two decades have been a major stimulus to renewed investi
gations into the origin and phylogeny of Primates. Moreover, assessments of
possible tupaiid-primate affinities have focused attention on the wide range of
molecular and soft anatomical evidence that can be used to supplement, rather
than replace, the more traditional dental and skeletal data for phylogenetic
reconstruction.
On behalf of the contributors, I wish to thank Mr. Kirk Jensen for his
encouragement and continued support of this project. Special thanks are also
due to Miss Mary Markytan and Mrs. Edith Witt for their invaluable assistance
in preparing this volume for publication. This book was composed by Western
ix
PREFACE
Typesetting Company, Kansas City, Missouri, and the editor acknowledges the
assistance of Mr. Jerry Germany and Mr. Alex Kreicbergs in completing this
phase of the project.
w.P.L.
