Table Of ContentEvolutionary Relationships
among Rodents
A Multidisciplinary Analysis
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Volume 92-Evolutionary Relationships among Rodents: A
Multidisciplinary Analysis
edited by W. Patrick Luckett and Jean-Louis Hartenberger
Series A: Life Sciences
Evolutionary Relationships
among Rodents
A Multidisciplinary Analysis
Edited by
W. Patrick Luckett
School of Medicine
University of Puerto Rico
San Juan, Puerto Rico
and
Jean-Louis Hartenberger
Institute of Evolutionary Sciences
University of Montpellier II
Montpellier, France
Springer Science+Business Media, LLC
Proceedings of a NATO Advanced Research Workshop on
Multidisciplinary Analysis of Evolutionary Relationships among Rodents,
held July 2-6, 1984, under the auspices of NATO and Centre
National de la Recherche Scientifique in Paris, France
Library of Congress Cataloging in Publication Data
NATO Advanced Research Workshop on Multidisciplinary Analysis of Evolu-
tionary Relationships among Rodents (1984: Paris, France)
Evolutionary relationships among rodents.
(NATO ASI series. Series A, Life sciences; v. 92)
"Proceedings of a NATO Advanced Research Workshop on Multidisciplinary
Analysis of Evolutionary Relationships among Rodents held July 2-6,1984, in Paris,
France"—T.p. verso.
"Published in cooperation with NATO Scientific Affairs Division."
Bibliography: p.
Includes index.
1. Rodents—Evolution—Congresses. 2. Mammals—Evolution—Congresses. I.
Luckett, W. Patrick (Winter Patrick) II. Hartenberger, Jean-Louis. III. North Atlantic
Treaty Organization. Scientific Affairs Division. IV. Title. V. Series.
QL737.R6N25 1984 599.32'30438 85-9480
ISBN 978-1-4899-0541-3
ISBN 978-1-4899-0541-3 ISBN 978-1-4899-0539-0 (eBook)
DOI 10.1007/978-1-4899-0539-0
©1985 Springer Science+Business Media New York
Originally published by Plenum Press, New York in 1985
Softcover reprint of the hardcover 1st edition 1985
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
For more than 40 years, contributions to the knowledge of rodent evolution by Pro
fessors Rene Lavocat and Albert E. Wood have dominated and enriched this
scientific field. All of the speakers and discussants at the Paris Rodent Evolution
Symposium have had the opportunity to read papers by these investigators, and
to exchange ideas with them. In addition, their friendship and warmth are well
known to all of us. Thus, it is with great pleasure and respect that we dedicate
this volume to these two outstanding scientists and friends.
Rene lavocat Albert E. Wood
PREFACE
The order Rodentia is the most abundant and successful group
of mammals, and it has been a focal point of attention for compar
ative and evolutionary biologists for many years. In addition,
rodents are the most commonly used experimental mammals for bio
medical research, and they have played a central role in investi
gations of the genetic and molecular mechanisms of speciation in
mammals. During recent decades, a tremendous amount of new data
from various aspects of the biology of living and fossil rodents
has been accumulated by specialists from different disciplines,
ranging from molecular biology to paleontology. Paradoxically,
our understanding of the possible evolutionary relationships among
different rodent families, as well as the possible affinities of
rodents with other eutherian mammals, has not kept pace with this
information "explosion." This abundance of new biological data
has not been incorporated into a broad synthesis of rodent phylo
geny, in part because of the difficulty for any single student of
rodent evolution to evaluate the phylogenetic significance of new
findings from such diverse disciplines as paleontology, embryology,
comparative anatomy, molecular biology, and cytogenetics.
The origin and subsequent radiation of the order Rodentia were
based primarily on the acquisition of a key character complex:
specializations of the incisors, cheek teeth, and associated mus
culoskeletal features of the jaws and skull for gnawing and chewing.
Consequently, the most commonly used characters for studying rodent
systematics have been components of a single functional system -
the masticatory apparatus. Moreover, it is commonly believed that
major changes in the teeth and skull for increased efficiency of
the masticatory apparatus have evolved in parallel several times
within unrelated higher taxonomic groups of rodents. This accep
tance of widespread parallelism has inhibited assessments of phylo
genetic relationships among rodent higher taxa during the past 30
years. Indeed, it is commonly believed that the traditionally
defined major groups of rodents (Sciuromorpha, Myomorpha, Hystrico
morpha, and Protrogomorpha) represent grade levels of organization,
rather than monophyletic clades, and that many of the shared simi
larities of each grade (such as hystricomorphy) have been attained
independently by some of its members.
vii
viii PREFACE
Several factors have contributed to the disagreements that
surround analyses of rodent evolution. (1) The diversity of the
group is a consequence of its extensive evolutionary radiation
during Eocene-Oligocene times, and the fossil record for these
epochs remains incomplete in many important aspects. (2) The
extensive number of species and families of rodents makes it diffi
cult to study and evaluate any character complex in most or all
rodent taxa. (3) Much of the study of rodent systematics has
focused on a search for key characters that can be used in classi
fications, rather than an assessment of the biological basis for
shared similarities among different groups. (4) When evaluating
phylogenetic relationships among rodents, many investigators have
failed to distinguish among shared primitive retentions, shared
derived traits, and shared similarities that may be due to parallel
or convergent evolution. (5) Emphasis has been placed on the dis
covery of ancestor-descendant relationships during rodent phylogeny,
and less attention has been devoted to the assessment of sister
group relationships among taxa. Hypotheses of ancestor-descendant
relationship are restricted to studies of the incomplete and frag
mentary fossil record, whereas assessments of sister-group relation
ships can utilize observations from both fossil and living forms.
In recent years, there has been a renewed interest in the evo
lutionary analysis of mammalian relationships, related in part to
an increase in the kinds of data that are being evaluated (soft
anatomical, embryological, molecular, and chromosomal, as well as
the more traditional dental and cranioskeletal features). This
resurgence has also profited by the increased emphasis and consider
ation given to problems of character analysis and the assessment of
homology and convergence. As yet, most of this "new" (non-dental
or cranial) data has been presented in isolation from other character
analyses; consequently, we have evolutionary "trees" or classifi
cations based on molecular, chromosomal, or dental traits, with
little or no attempt to integrate these findings into a broader
assessment of rodent (or mammalian) phylogeny.
As an initial step toward evaluating and integrating a vast
amount of biological data into an analysis of evolutionary rela
tionships among rodents, we organized a NATO Advanced Research
Workshop that was held in Paris, France during July 2-6, 1984.
This symposium provided a forum for a broad exchange of views among
specialists from the different disciplines of paleontology, compar
ative anatomy, embryology, molecular biology, and cytogenetics, and
this volume is the proceedings of that symposium. Participants
were asked to discuss their methodology of phylogenetic reconstruc
tion, and, in particular, their criteria for evaluating homology,
convergence, and the relatively primitive or derived nature of
shared similarities used in evolutionary analyses. The focal point
of the symposium was on character analysis and phylogenetic recon
struction from a wide range of data, and the use of these data to
PREFACE
test hypotheses of ancestor-descendant and cladistic relationships,
both within the order Rodentia and between rodents and other
eutherian orders. The initial portion of the volume is devoted
to an assessment of the eutherian affinities of the Rodentia, and
later sections focus on specific questions of intraordinal relation
ships at the family level or above.
We have concentrated on the multidisciplinary analyses of evo
lutionary relationships among rodents, and we have avoided the temp
tation of proposing new classifications of rodents, based on our
preliminary analyses. Several hypotheses of higher taxon phylogeny
were corroborated by analyses in this volume, including the mono
phyly of the Hystricognathi and Muroidea. Other problems of sys
tematic relationships remain to be clarified, and it is our hope
that future investigations will attempt to resolve these contro
versies by multidisciplinary analyses. If future research on rodent
evolution is accomplished by collaborative efforts between paleon
tologists and neontologists, then one of the main goals of this
symposium and volume will have been attained.
The editors are indebted to the following organizations for
their financial support for the Paris Symposium, and for the sub
sequent preparation of this volume: North Atlantic Treaty Organi
zation (Scientific Affairs Division); Centre National de la Recherche
Scientifique (secteurs Sciences de la Vie et Terre, Ocean, Atmo
sphere, Espace); Ministere de l'Industrie et de la Recherche (Fonds
a
d'aide la recherche No. 84.C.0431); and the National Science
Foundation. We wish to offer our special thanks to Madame Christiane
Denys for her administrative assistance during the organization of
the Paris conference. We acknowledge and appreciate the scientific
advice of our colleagues Drs. Jean Chaline, Rene Lavocat, and John
Wahlert during the initial phases of organizing the symposium.
Technical assistance for the symposium and volume was provided by
Madame Genevieve Jean, Monsieur Serge Legendre, and Mademoiselle
Elisabeth Natale in Montpellier, and Senora Ivette Melendez in San
Juan. Various aspects of editorial assistance in preparing this
volume were furnished by Ms. Pat Vann and Mr. John Matzka at Plenum
Press, and we thank them for their efforts.
A special debt of gratitude is owed to Dr. Nancy Hong for her
extensive help in all phases of the organization of the Paris
Symposium, as well as for her aid in the preparation and editing
of this volume. This project could not have been produced without
her tireless and unselfish assistance.
W. P. Luckett
J.-L. Hartenberger
CONTENTS
ORIGIN AND EUTHERIAN AFFINITIES OF RODENTIA
The Order Rodentia: Major Questions on Their
Evolutionary Origin, Relationships
and Suprafamilial Systematics • • • • • 1
J.-L. Hartenberger
Possible Phylogenetic Relationship of Asiatic
Eurymylids and Rodents, with Comments
on Mimotonids • • • • • • • • • • 35
C.-K. Li and S.-Y. Ting
Cranial Evidence for Rodent Affinities 59
M. J. Novacek
Rodent and Lagomorph Morphotype Adaptations,
Origins, and Relationships: Some
Postcranial Attributes Analyzed • • • • 83
F. S. Szalay
Enamel Structure of Early Mammals and Its Role in
Evaluating Relationships among Rodents • • • • • • 133
A. Sahni
Reconstruction of Ancestral Cranioskeletal
Features in the Order Lagomorpha • • • • • • • • • 151
N. Lopez Martinez
A Phylogeny of Rodentia and Other Eutherian Orders:
Parsimony Analysis Utilizing Amino Acid
Sequences of Alpha and Beta Hemoglobin Chains ••••• 191
J. Shoshani, M. Goodman, J. Czelusniak, and
G. Braunitzer
