Table Of ContentTHE 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
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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