Evolutionary Relationships among Rodents: A Multidisciplinary Analysis PDF
Preview Evolutionary Relationships among Rodents: A Multidisciplinary Analysis
Evolutionary Relationships among Rodents A Multidisciplinary Analysis NATO ASI Series Advanced Science Institutes Series A series presenting the results of activities sponsored by the NA TO Science Committee, which aims at the dissemination of advanced scientific and technological knowledge, with a view to strengthening links between scientific communities. The series is published by an international board of publishers in conjunction with the NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics New York and London C Mathematical D. Reidel Publishing Company and Physical Sciences Dordrecht, Boston, and Lancaster D Behavioral and Social Sciences Martinus Nijhoff Publishers E Engineering and The Hague, Boston, and Lancaster Materials Sciences F Computer and Systems Sciences Springer-Verlag G Ecological Sciences Berlin, Heidelberg, New York, and Tokyo Recent Volumes in this Series Volume 86-Wheat Growth and Modelling edited by W. Day and R. K. 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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
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