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292 Pages·1995·9.031 MB·English
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THE EVOLUTIONARY ECOLOGY OF ANIMALS STUDIES IN SOVIET SCIENCE LIFE SCIENCES 1973 MOTILE MUSCLE AND CELL MODELS N. I. Arranet PATHOLOGICAL EFFECTS OF RADIO WAVES M. S. Talgskaya and Z. V. Gordan CENTRAL REGULATION OF THE PITUITARY-ADRENAL COMPLEX E. V. Naumenka 1974 SULFHYDRYL AND DISULFIDE GROUPS OF PROTEINS Yu. M. Tarchinskii MECHANISMS OF GENETIC RECOMBINATION V. V. Kushev 1975 THYROID HORMONES: Biosynthesis, Physiological Effects, and Mechanisms of Action Ya. Kh. Turakulav, A. I. Gagergans, N. S. Salakhava, A. K. Mirakhmedav, L. M. Garber, V. I. Kandrar, and G. A. Gaidina 1977 THE EVOLUTIONARY ECOLOGY OF ANIMALS S. S. Shvarts HEMATOPOIETIC AND LYMPHOID TISSUE IN CULTURE E. A. Luriya STRUCTURE AND BIOSYNTHESIS OF ANTIBODIES R. S. Nezlin PROTEIN METABOLISM OF THE BRAIN A. V. Palladin, Ya. V. Belik, and N. M. Palyakava 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. STUDIES IN SOVIET SCIENCE THE EVOLUTIONARY ECOLOGY OF ANIMALS S. S. Shvarts Academy of Sciences of the USSR, Urals Branch Sverdlovsk, USSR Translated from Russian and edited by Ayesha E. Gill University of California, Los Angeles With new material by the author and editor CONSULTANTS BUREAU· NEW YORK AND LONDON Library of Congress Cataloging in Publication Data ShvarfS, Stanislav Semenovich. The evolutionary ecology of animals. (Studies in Soviet science) Translation of EvolillfSionnalcl ekologilcl zhivotnykh. Includes bibliographies and index. 1. Evolution. 2. Zoology-Ecology. 3. Population genetics. I. Title. II. Series. QH371.S4813 575 76-50647 ISBN 978-1-4684-8099-3 ISBN 978-1-4684-8097-9 (eBook) 001 10.1007/978-1-4684-8097-9 The original Russian text of The Evolutionary Ecology of Animals was published by the Urals Branch of the Academy of Sciences of the USSR in Sverdlovsk as Vol ume 65 of the Proceedings (Trudy) of the Institute of Plant and Animal Ecology in 1969. It was corrected by the author for the present edition. This translation is published under an agreement with the Copyright Agency of the USSR {VAAPl. 3B0J1IOUI10HHA5I 3K0J10rJ.151 )KI1BOTHblX C. C. WBAPU EVOLYUTSIONNAYA EKOLOGIYA ZHIVOTNYKH S. S. Shvarts © 1977 Consultants Bureau, New York Softcover reprint of the hardcover 1s t edition 1977 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 p'ermission from the Publisher Editor's Foreword While translating this book, I was in close communication with the author, S. S. Shvarts (Schwarz), who read and commented on the entire translated manuscript. In particular, any ambiguities as to the identity of organisms described only by common names in the original text were removed, because the author kindly supplied the Latin names in all such cases. Com mon names are retained in the translation, but the Latin names are also added where needed. Some of the terminology used in the Russian is a transliteration from English words employed now more by European workers than Americans. I have defined these terms or noted their more common equivalents used in current American literature where it seemed useful in the text. A final chapter, "Recent Work on the Evolutionary Ecology of Ani mals," is presented as Appendix II to the translation of the original text. I have written this chapter in order to update the material presented in the original edition published in 1969. The chapter discusses important recent contributions relevant to the subject matter presented by Shvarts. I would like to thank W. Z. Lidicker, Jr., and Y. B. Linhart for reading this final chapter and providing very helpful suggestions and comments. I am particularly grateful to the author, S. S. Shvarts, for his careful reading of the translated manuscript. Ayesha E. Gill Los Angeles, California [Prof. Shvarts died May 12, 1976] v Contents Preface 3 Introduction 5 Chapter I. The Genetic Basis of the Reorganization of Populations 21 Chapter II. An Ecological Estimate of Interpopulation Differences 33 Chapter III. The Reorganization of Populations-Homeostatic Alterations of the Genetic Structure of Populations and Microevolution 49 Chapter IV. Ecological Mechanisms for the Maintenance of the Genetic Heterogeneity of a Population 75 Chapter V. Ecological Mechanisms for the Reorganization of the Genetic Structure of Populations 91 1. Role of the Dynamics of Population Age Structure in the Reorganization of Its Genetic Makeup 91 2. On the Significance of Nonselective Elimination 110 3. The Spatial Structure of Populations-A Factor in Microevolution 120 Chapter VI. Speciation 145 Chapter VII. The Ecological Essence of Macroevolution 193 Conclusions 215 References and Bibliography 223 Appendix I. The Species Problem and New Methods of Systematics 257 Appendix II. Recent Work on the Evolutionary Ecology of Animals (by A. E. Gill) 275 vii The Evolutionary Ecology of Animals Ecological Mechanisms of the Evolutionary Process S. S. Shvarts Preface Over a period of 20 years, the specific rules determining the ecological and morphophysiological features of populations of different species under dif ferent environmental conditions were studied in the Zoological Laboratories of the Institute of Biology, Urals Branch, of the Academy of Sciences of the USSR.* All classes of terrestrial vertebrates were studied. Research was conducted both under natural conditions in diverse biomes (from the step pes to the tundra) and experimentally on model populations. The material accumulated during this period led us to conclusions concerning the mechanisms of the evolutionary modifications of animals. Data were obtained that indicated the possibility of utilizing the animals' reactions in the alteration of environmental conditions during an elaboration of the most difficult problems of speciation. We presented the synthesis of these data in a monograph (Shvarts, 1959). Another series of investigations was devoted to the study of the dynamics of population structure. These investigations led us to the conclusion that alteration of the intrapopulation structure of a species leads to an alteration in the genetic makeup of intraspecific group ings-to microevolutionary modifications. The results of this work were published in several journals (Shvarts, J963a-c, 1965, and others). A general analysis of the data obtained in our laboratories along the two paths indicated made evident the deep inner connection between the mor phophysiological reactions of different forms and the alteration of popula tion structure. On this basis, the representation of the ecological mechanisms of the evolutionary process-the study of which constitutes, in our opinion, the main task of evolutionary ecology-matured. Synthesis of * Since 1965, the Laboratory of the Population Ecology of Vertebrates of the Institute of Plant and Animal Ecology. 3 4 THE EVOLUTIONARY ECOLOGY OF ANIMALS the results of morphophysiological and population-ecological investiga tions, however, required the study (both experimental and theoretical) of certain questions that until recently had not attracted our attention. The most important of these questions are the evolutionary significance of the dynamics of population spatial structure, the correlation of micro- and macroevolution, and the reality of taxa above the species level. The results of these investigations are published for the first time in this book. The basic task of the book is to present in a well-grounded way the key signifi cance of ecological mechanisms in the evolutionary process. We view these mechanisms, together with natural selection, as the basic motive force of evolution. Both the structure of the book as a whole and the structure of individual chapters are subordinate to this task. In particular, we use data from the literature only insofar as they are necessary for the objective analysis of the basic problems of evolutionary ecology. We did not strive to give a comprehensive summary of the literature on all the subjects broached. There is no necessity to do this, since a critical review of the literature on the evolutionary genetics of populations has been published by Dubinin (1966a). This book is, to a considerable degree, the theoretical result of the work of a large collective of scientific workers and laboratory assistants. Particularly great contributions to the general work, which I aspired to synthesize in this book, were made by V. N. Pavlinin, V. S. Smirnov, V. E. Bergovyi, V. N. Bol'shakov, L. N. Dobrinskii, Z. D. Epifantseva, V. G. Ishchenko, N. A. Ovchinnikova, V. G. Olenev, A. V. Pokrovskii, O. A. Pyastolova, L. M: Syuzyumova, and L. K. Yashkova. I deeply thank all my colleagues, collaborators from the Laboratory of the Population Ecology of Vertebrates, whose creative work made it possi ble to accumulate material for the statement of certain principal questions of evolutionary theory. Introduction The majority of contemporary biologists regard evolution as a process of progressive mastery of the arena of life by animals and plants, as progressive adaptation to diverse conditions of existence. In this respect, even the adherents of concepts of spontaneous generation (in any of their variants) represent no exception, since the result of spontaneous generation is, in the final analysis, evaluated and corrected in the process of the organism's interaction with its environment. Hence, it is clear that any evo lutionary conception broaches, to some degree or other; the subject of the interaction of an organism with its environment. It is natural, also, that any evolutionary theory is obliged to rest on ecological laws, for, irrespective of the well-known transformation of biologists' views on the purpose <ind method of ecology, its basic task has remained unchanged now for the course of a hundred years. This task is to investigate the lives of animals and plants in their natural habitats, in nature. If one speaks not of evolutionary theory in general, but of the only evo lutionary theory that has withstood the test of time and satisfied the demands of modern science-of Darwinism-then its ecological basis is clear. Petrusewicz (1959) is correct in entitling one of his theoretical works "Darwin's Theory of Evolution Is an Ecological Theory" [in Polish]. The author correctly emphasizes that evolution is in essence an ecological process, accomplished on the basis of laws governed by group, not indi vidual, phenomena. Orians (1962) is even more categorical, asserting that the general theory of ecology is a theory of natural selection. The attention paid by evolutionists to ecological rules has intensified especially in recent times in connection with the establishment of certain general laws of ecology, reflecting in its most general form the interrelation ship of organisms with their environment. On the other hand, the role of 5

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