Russian Studies in History

ISSN: 1061-1983 (Print) 1558-0881 (Online) Journal homepage: http://www.tandfonline.com/loi/mrsh20

From the History of Soviet Ecology

Eduard N. Mirzoian

To cite this article: Eduard N. Mirzoian (1995) From the History of Soviet Ecology, Russian
Studies in History, 34:2, 7-23

To link to this article: http://dx.doi.org/10.2753/RSH1061-198334027

Published online: 09 Dec 2014.

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 EDUARD
N. MIRZOIAN

From the History of Soviet Ecology

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The Evolutionary Views of D.N. Kashkarov

For centuries, natural scientists have emphasized the role of the environ-
ment in humanity’s destiny [ 13. In our own day, the world community
has finally realized that its common future is inseparably linked to the
preservation of the biosphere [2]. Throughout its history, civilization
has faced the necessity of assuming control of global processes,
which constitute the foundation of vital activity and the evolution of the
biosphere. One of the first to make this point was V.N. Beklemishev,
but this eminent biocenologist’s warning was persistently ignored [3].
In order to solve a problem of such magnitude as the management of
the biosphere, natural science must mobilize all its resources, and in
particular, of course, reconsider ecology from this viewpoint.
Research on the history of ecology, in particular Soviet ecology,
should be an inseparable component of that general effort. It should
not be forgotten that in our country the process of deformation affected
fields other than genetics. The criticism of genetics became a launch-
ing pad for a much broader assault. A blow was struck against all
theoretical biology, and in particular its very heart-evolutionary biol-
ogy. First-nd under the banner of defending Darwinism, too!-T.D.
Lysenko waged a struggle against N.I. Vavilov, who had achieved a
synthesis of the theory of evolution and genetics in the interests of

Russian text 0 1991 by E.N. Mirzoian. “Iz istorii otechestvennoi ekologii:

evoliutsionnye vozzreniia D.N.Kashkarova,” Voprosy istorii estestvoznuniiu i
tekhniki, 1991, no. 4, pp. 13-21. A publication of the Institute of the History of
Science and Technology, Russian Academy of Sciences.
Eduard Nikolaevich Mirzoian, Doctor of Biological Sciences, is the head of the
Sector for the History of Biology, Institute of the History of Science and Technol-
ogy, Russian Academy of Sciences.
7
 8 RUSSIAN STUDIES IN HISTORY

both biology and selection [selektsiia] [4]. Then Lysenko discredited

Darwinism itself, calling it a passive and trivial evolutionary doctrine.
A broad new evolutionary synthesis presented by Academician 1.1.
Shmal’gauzen in his Problems of Darwinism [Problemy Darvinizma]
(1946) was subjected to scathing criticism at the August 1948 session
of VASKhNIL [V.I. Lenin All-Union Academy of Agricultural Sci-
ences]. Soon afterward, a savage assault was launched against the
teachings of Academician A.N. Severtsov, who had developed a mod-
em theory of the macroevolution of the organic world. In 1950, a joint
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session of the USSR Academy of Sciences and the USSR Academy of

Medical Sciences struck the leader of evolutionary physiology, Acade-
mician L.A. Orbeli, with a crippling blow, and the research directions
that he had been developing were drastically impeded (delayed). Criti-
cism was also directed against the leader of evolutionary paleontology,
Academician A.A. Borisiak. Even earlier, from the 1930s onward, un-
justified criticism was directed against the founder of evolutionary
histology, Academician A.A. Zavarzin, and a theory he had developed
concerning the laws governing the evolution of tissues became a target
of attack. Two major centers of evolutionary thought, headed by N.K.
Kol’tsov and S.S. Chetverikov, ceased to exist. Ecology was the target
of not very competent criticism over a span of several decades. In the
1920s and 1930s, accusations of serious philosophical errors were di-
rected against the country’s leading phytocenologists and biocenologists:
I.K. Pachosskii, L.G. Ramenskii, A.P. Il‘inskii, V.V. Alekhin, V.N.
Sukachev, D.N. Kashkarov, and V.N. Beklemishev. In the 1930s the
personal intervention of T.D. Lysenko and 1.1. Prezent put an end to
the development of a fruitful line of research in biocenology developed
by V.V. Stanchinskii, which tragically affected the fate of this talented
scientist. In the early 19.50s Lysenko made an attempt to denigrate
biogeocenology and its acknowledged leader, Academician V.N. Suka-
chev. Both Sukachev and Stanchinskii had invested considerable effort
in synthesizing biocenology with the theory of evolution, the principles
of genetics, and V.I. Vemadskii’s theory of the biosphere. In 1951 a
conference of ecologists denounced several scientists who had made
contributions to evolutionary ecology [5]; D.N. Kashkarov was among
those attacked. Along with G.F. Gauze, A.N. Formozov, and N.I. Kala-
bukov he was denounced as an opponent of “Michurinist ecology.” As
the aforementioned ecological forum passed its “resolution,” D.N.
Kashkarov, however, was no longer among the living; he died in
 FALL 1995 9

November 1941 at Khvoinaia Station during the evacuation from block-

aded Leningrad.a
Daniil Nikolaevich Kashkarov played a prominent role in the devel-
opment of ecology in the USSR. A widely educated and versatile
scientist, who skillfully combined field research with experimentation
and the systematic gathering of factual material with theoretical gener-
alization, Kashkarov left a profound stamp on the creation and devel-
opment of evolutionary ecology. His evolutionary views constitute a
part of his overall ecological conceptions and deserve to be examined
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separately.
The development of modem ecology in our country is linked to
Kashkarov’s name [6]. He began his scientific career as a morpholo-
gist. While still a college student under the influence of M.A. Menzbir,
he chose zoology as his specialty and began working in his teacher’s
laboratory, researching a topic in comparative anatomy. During his
student years Kashkarov was also greatly influenced by V.A. Vagner,
who awakened in him an interest in zoopsychology. Kashkarov’s earli-
est zoological works already reveal his interest in problems of ecology.
After 1920, when he moved to Tashkent and organized the Turkestan
(later the Central Asian) State University, ecology became his life’s
work. In 1924 he began to teach a course of lectures on the ecology of
animals. The shift in focus of Kashkarov’s scientific interests is natu-
rally tied to his affiliation with the school of zoologists at Moscow
University, in which an ecological tradition had existed since the time
of K.F. Rul’e [7]. Kashkarov’s thorough, general biological training,
his profound knowledge of Darwinism and his understanding of its
ecological orientation, and his striving for broad theoretical generaliza-
tions made him decide, after he had settled in Tashkent, to devote
himself to ecology, which would provide connections to various divi-
sions of biology, to the theory of evolution, and to practical work. In
1928, Kashkarov undertook a research visit to the United States, where
he became acquainted with the works of American ecologists such as
Charles [Christopher] Adams, Victor Shelford, Frank Chapman,
George Grinnell, U. Taylor [name transliterated], and Charles Voorhis.
“After his trip to the United States, D.N. [Kashkarov] moved entirely
into the field of ecology, which remained his very favorite discipline to
the end of his life” [8, p. 711. In 1933, Kashkarov was elected to head
the department of vertebrate zoology at Leningrad State University.
There he organized an animal ecology laboratory. He founded an ecology
 10 RUSSIAN STUDIES IN HISTORY

committee attached to the Leningrad Society of Natural Scientists, and

a journal, V o p r o v ekologii i biotsenologii [Issues in Ecology and Bio-
cenology],* which began to appear in 1934 (only seven issues were
published). Kashkarov supervised the organization of the USSR’s first
All-Union Ecology Conference in Leningrad in 1938.
Having decided to work in ecology, Kashkarov refused simply to
copy existing models. He thought out for himself the factual material
and the theoretical principles of ecology. In setting up his ecological
platform he drew upon the theory of evolution and on the ecological
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tradition that he had inherited from K.F. Rul’e, K.M. Ber, A.F. Mid-
dendorf, N.A. Severtsov, and V.O. Kovalevskii. Kashkarov received
this tradition from his teacher, M.A. Menzbir. His friend and coauthor
V.V. Stanchinskii underlined the importance of this tradition. Although
he gave due credit to his predecessors’ ecological ideas, Kashkarov
nonetheless believed that ecology was still in its formative stage, and
that its subject, tasks, and methods were still in the discussion stage.
To understand better ecology’s place and tasks in the system of biolog-
ical sciences he turned to the history of biology. He divided the post-
Darwinian period of biology’s development into two stages. The first
stage was marked by a fascination with the task of reconstructing the
paths of phylogenesis and by a growth in experimental research. The
whole organism, functioning in a particular environment, was forgot-
ten; in their striving to explain organisms’ lives, scientists frequently
“studied only isolated parts of them” [9, p. 321. These contradictions
were overcome during the next stage, marked by a period of “striving
for synthesis, a period of combining various divisions of biology into
one” [ibid.]. Enriched by the achievements of the analytic period, biol-
ogy “returned to the natural world” [ibid.]. Kashkarov clearly realized
that the 1920s and 1930s were to play a decisive role in biology’s fate.
He gave considerable thought to ways to resolve the confrontation
between “classical” and “experimental” biology, which path the theory
of evolution should take, and, most important, what place ecology was
to occupy in these events.
Kashkarov had a very broad understanding of ecology’s purpose
and its place in the system of sciences; he viewed it as “a separate
discipline, a special orientation, a particular approach to things” [ 10, p.
1301, and even as a “method of thinking” [ 11, p. 371. Defining ecology

*Successor to Zhurnal ekologii i biotsenologii (1931). -Ed.

 FALL 1995 11

as the science of relations between an organism and the physical fac-

tors of the environment,or as the science of adaptations, did not satisfy
him. “Ecology is the science that studies the reactions of organisms
(both individual species and groups of organisms which are known as
communities or biocenoses) to the environment around them, reactions
that for the most part consist of adapting to their native habitat” [ 12, p.
71. Kashkarov did not share the notion that animal ecology was an
offshoot of zoogeography. He believed that the birth of ecology had
been stimulated by evolutionary theory.
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The biogeographer has no interest in problems of adaptation or the

philosophy of that process; he is not interested in the process of species-
formation in and of itself,or the role of the environmentin that process,
the role of physiology and history, of struggle and natural selection. . . .
He is not interested in the life or structure of communities, or the
dynamics of their life in connection with the dynamics of the environ-
ment. Yet all of these problems constitute the foundation of ecology
112, p. 153.

The necessity of studying all these processes in nature gave birth to

ecology.
Hence, according to Kashkarov, the theory of evolution constitutes
the foundation of ecology, and the range of problems of evolution
forms part of the task of ecological research. Autecology, synecology,
and biocenology must interact in studying the evolutionary process. In
response to the question of just what an ecologist is in a position to
study, Kashkarov answered: “Every phase of the evolutionary process,
so that we may get a total picture of it” [ 12, p. 2021. In his exploration
of this scheme he took the ecological point of view in his examination
of mutability, the struggle for existence, the effect of natural selection,
and species-formation.Later he expanded the sphere of possibilities of
evolutionary ecological research, and found that “ecology can play a
major role in the study of both the processes of species evolution and
the evolution of the biological community (biocenoses)” [9, p. 3141.
His own works and the experience of world science enabled Kashka-
rov to conclude that ecology can serve the evolutionist as a powerful
tool in reconstructing the paths of evolution and the laws that govern it
[ibid.]. He asserted, moreover, that the ecologist’s path to this set of
problems lay through the mastery of the latest advances in biology. It
is no accident that as he explored the sources of biogeocenology and
 12 RUSSIAN STUDIES IN HISTORY

evolutionary ecology in our country he emphasized: “We are turning

again to the natural world, but we are returning with a different ap-
proach, enriched by advances in the historical method, armed with
skills created in the laboratory and with thinking that has been disci-
plined by comparative morphology” [ 10, p. 1171.
In mapping out his program of ecological research, Kashkmv turned
to the theory of evolution in order to evaluate the situation in evolu-
tionary biology and the role of ecology in a new, maturing evolution-
ary synthesis. He determined that after Darwin’s The Origin of Species
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was published, “people based their thinking on the concept of struggle

for existence but did not gather factual material” [ 12, p. 2031. Yet only
a profound understanding of the interrelationships of species and indi-
viduals, as well as organisms and the environment, provided the pre-
requisite for the study of the effect of natural selection in nature. “It is
impossible, of course, to prove the presence of selection only because
an immeasurably larger number of individuals come into the world
than survive. It is essential to go out into the field for facts and ideas”
[12, p. 2051. To Kashkarov’s way of thinking, all the fundamental
postulates of Darwinism needed to be reinvestigated. And he assigned
a crucial role, in this monumental undertaking, to genetics and ecol-
ogy. As he, like other evolutionists of the 1930s, gave thought to a
strategy for a new evolutionary synthesis, he concluded that in con-
structing a theory of species-formation “the geneticist and ecologist
must work hand in hand,” that the results of their researches in that
field would “coincide with one another,” and that the processes of
species-formation could “not be understood without the aid of ecol-
ogy” [12, p. 1981.
Kashkarov returned to this problem again and again. As he explored
the logic of the development of the theory of evolution he asserted:
at the present stage of development of evolutionary theory, no other
biological discipline contributes as much to that theory as ecology does,
united as it is with genetics to study the process of species-formation
(small systematic units), and studying as it does the most fundamental
problem of ecology-the question of the relationship between the or-
ganism and its natural habitat, having to do with the community, the
struggle for existence, selection, and so forth I: 13, p. 181.
While he advocated an alliance between ecology and genetics, he
nevertheless saw a precise demarcation between the areas of interest of
 FALL. 1995 13

the two disciplines. “Evolution has originated and proceeds under the
control of two basic factors: heredity and the environment. Genetics
deals primarily with the former, while ecology deals mainly with the
latter. And the evolutionist must never neglect ecology or fail to apply
the ecological method of investigation” [ 14, p. 4931. The definition of
the environment here includes both its physical conditions and the
biotic environment-that is, “the community in which the evolving
organisms have lived and functioned” [ibid.].
During the 1920s and 1930s, all ecologists did not include problems
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of evolution within their science. Sometimes, polemics erupted be-

cause of this. Thus, for example, Kashkarov wrote: “Contrary to Shel-
ford’s opinion, the ecologist can and must be concerned with questions
of evolution” [14, p. 5141. The range of these issues was vast, and
Kashkarov assigned to it specific questions such as mutability, the
reproductive rate of individuals of a species, migration, the struggle for
existence, the adaptivity of species characteristics, the role of isolation
and species-formation, natural selection, and phylogeny. At the foun-
dation of evolutionary ecology he placed the evolutionary process,
which he had created by synthesizing genetic and ecological concep-
tions. According to this scheme, a species manifests variable phenotyp-
ical mutability, and under conditions that are normal to it a range of
deviations from the average may be observed. This mutability is gener-
ated by differences both among the organisms and among factors of
the environment. Changes in the phenotype are of an adaptive charac-
ter, but they are not inherited; “the genotype remains untouched” [14,
p. 4961. Competition arises among the altered individuals. Factors that
lie near the boundary of the organism’s possibilities, in the minimum
or maximum position, determine the selection of individuals; this, how-
ever, “does not lead to a change in the species” [ibid.]. A different
result will come about if particular factors affect individuals with a
force that exceeds the norm. External factors can cause mutations that
proceed directionally [napravfenno]. In that case, “the progeny of these
individuals will manifest gene variations, a bouquet of gene variations
in a great variety of directions” [ibid.]. Among these may be lethal,
harmful, neutral, and beneficial variations. Moreover, “some of the
mutated individuals will remain completely viable (Muller, Timofeev-
Resovskii . . .). From this diversity of mutations, which have been
caused by the fact that not all gametes or embryos are identical,. . .
selection may extract adaptive ones and destroy those that are not in
 14 RUSSIAN STUDIES IN HISTORY

tune with the environment” [9, p. 2791. In a few cases, many individu-
als will change in one direction, thus creating the prerequisites for the
development of new life-forms ~omoobruzovunie].
Any one of the new gene variations may end by becoming established
on the spot, or elsewhere thanks to dispersal under especially favorable
conditions, and continue to exist and produce progeny which inherits its
features, which will vary around an average. In the struggle for life, in
the process of competition, the bouquet of new gene variations may
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subsequently produce not one but two, three, or more new genotypes.
And then we have the divergence of one species into several new ones
[ 14, p. 4963.

The fate of the new forms will ultimately be decided by ecological

factors. Forms that originate randomly may be preserved and fit into a
given biotype only if their properties and features prove to be in con-
formity with the environment they inhabit. To Kashkarov’s way of
thinking, the new, small systematic units into which the varying spe-
cies fall ought to be the object of ecology. “A thorough, detailed study
of them in connection with the circumstances under which they live
and function-an ecological study-can contribute a great deal to our
understanding of species-formation and of the role of the environment
in that process” [ 14, p. 4981.
Kashkarov paid special attention to the initial stages of species-
formation. “What is it that compels a species to create new forms and
to divide up into ecological races?” [14, p. 5131 he asked. There were
various answers to this question. Kashkarov, however, assigned to eco-
logical factors the role of the starting mechanism in species-formation.
“The emergence of new races most often starts with a change in the
ecological optimum: the choice of a new habitat entails a change in
reactions to physical factors, which are then followed by the acquisi-
tion of morphologically adaptive features under the influence and the
control of the new conditions” [14, p. 5141. The choice of a new
habitat also means overcoming a certain barrier. Just how does a spe-
cies manage that? “The overcoming of the ecological bamer and, as a
result, the expansion of the areal range of a species (whether ecological
or geographical) take place because in the different contacting biotypes
or zones, new forms come into being (ecotypes, races, varieties), forms
that are distinct from one another ecologically and sometimes in form
as well” [ibid.].
 FALL 1995 15

Kashkarov ’s understanding of the process of species-formationtook

shape under the remarkable influence of new views on the Linnaean
species as a system, views developed in our country by N.I. Vavilov
and his school. In accordance with these views, Kashkmv acquired a
concept of species as a community of biotypes united by the history of
their origin, and a concept of an ecotype as a group of biotypes of one
species. With corrections for the most recent ecological concepts, the
pattern of species-formation looked like this: “Evolution is adaptive; a
species splits up into small, systematic units because of the effect of
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natural selection, which, within one species, creates a group of bio-

types that are in conformity with particular climatic, edaphic, or biotic
conditions of the habitat” [15, p. 6231.
In mapping out approaches to the study of species-formation via
ecology, Kashkarov made extensive use of the experience of the N.I.
Vavilov school and the results of V.V. Stanchinskii’s research. Having
remarked that zooecology was beginning to accumulate material attest-
ing that the process of formation of ecotypes takes place in the animal
world as well, Kashkarov emphasized that “the study of ecotypes is of
enormous theoretical interest, because it leads us directly to the study
of the process of species-formation, the study of the process of evolu-
tion against the background of the environment” [9, p. 2971. Data
obtained from investigating the fauna of deserts convinced him that as
a rule, a species does not leave the boundaries of its own area, because
conditions suitable for it are lacking in neighboring biotypes. If a spe-
cies turns out to be able to occupy other biotypes besides its own, it
changes its habits and its reactions to the environment. “This marks the
beginning of the formation of a new species” [9, p. 3003.
The next stage in species-formation is a change in structure, which
follows a change in habit. The assertion of such a sequence of events
had nothing in common with Lamarckism. Kashkarov based his work
on the principles of population genetics. “What we have here is a
certain sequence of development which is the result of the environment
and the selection of features from innumerable gene variations,” he
wrote, “a sequence conditioned by the fact that the emergence of new
features in a structure in the absence of a move to a new habitat where
the feature might prove to be adaptive-that is, without a change in
behavior-will not be advantageous to the species” [ibid.]. Kashkmv
insisted that the problem of the adaptivity of species features and their
role in species-formation belonged in the realm of evolutionary ecology.
 16 RUSSIAN STUDIES IN HISTORY

While Charles Darwin attributed adaptive significance not only to spe-

cies features but also to features of smaller systematic units, later re-
searchers often attributed adaptive significance only to features of the
major systematic categories such as classes, orders, and families. Hence,
“the Darwinian idea of the adaptivity of evolution seemingly collapsed”
[9, p. 2933. Kashkarov did not accept notions of this sort. He believed
that the real species features were those that “give a species an advan-
tage in the struggle for existence and make it the victor in the struggle
of life, enabling it to populate new zones and allowing it to take pos-
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session of territory that another species could not possess, or enabling

two species living side by side not to fight one another but each to
occupy its own niche in the community of nature” [9, p. 2951. These
are the adaptive features. They are rarely expressed morphologically;
they are mostly physiological and etiological features which find ex-
pression “in the behavior of the species and its reactions to particular
conditions, in its relation to physical factors of the environment, in its
choice of habitat, places of reproduction, feeding, and so forth” [ibid.].
Morphological features take on adaptive significance gradually. At
first, morphological differences may be lacking altogether; afterward,
they are expressed in very small ways and only later still “become ever
larger and, subjected to selection, also adaptive” [9, p. 3091. In this
way, according to Kashkarov, “the higher the systematic categories
that we deal with, the more adaptive their distinguishing features be-
come” [ibid.]. Such an interpretation of the process of the emergence
of adaptive features opened the way to ascribe more active involve-
ment to ecology in constructing the theory of species-formation and
macroevolution.
During the 1920s and 1930s, no one discipline could claim single-
handedly to have solved the problem of species-formation. People in-
creasingly become convinced that the research being conducted in the
various fields-in particular, genetics and ecology+xght to comple-
ment one another. However, this conviction was not shared by every-
one; a number of scientists believed that any appeal to the environment
would deflect the study of evolution from the path laid down by genet-
ics. To this, Kashkarov objected: “It is a profound error to think that
the study of ecology leads to Lamarckism. Moreover, in matters of
species-formation, it is impossible to gain understanding without the
aid of ecology” [14, p. 5181.
The role attributed to ecology increased still more during the study
 FALL 1995 I7

of evolution above the species level. Data from paleozoology led re-
searchers to think that “bursts in the evolutionary process occurred in
connection with major changes in the earth’s orographic and climatic
state” [ibid.]. In exactly the same way, observations of modem animals
led to the conclusion that “the evolutionary process is inseparably
linked to changes in the environment” [ibid.]. Kashkarov refined and
elaborated upon this overall postulate and concluded that both species-
formation and the evolution of taxons are inseparably linked to the
evolution of ecosystems. He expressed the unity and the intercon-
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nection of all three of these components of the evolutionary process in

the form of several postulates that taken together form a theory of the
ecological laws of evolution. It is based on Darwin’s postulate of
the link between species-formation and the conditions of abiotic and
biotic environments. Kashkarov confirmed that adaptation and evolu-
tion, on the level of species, take place “in very close relation to the
environment” [ 14, p. 5201. He went on to show that the evolution of a
species takes place “in close correlation with changes in other organisms
which make up the natural community” [ibid.]. What was new in this
was the idea of the biotic environment as an organized ecosystem.
Kashkarov insisted that a biocenosis is not merely a mechanical aggre-
gate of species and that selection on the level of ecosystems does not
simply enable species to adjust to one another, but that it serves to
foster the organization of the ecosystem as a whole. The organization
and integrity of ecosystems are manifested in their vital activity. “A
biocenosis is a very complex system in which the individual compo-
nents are not only compatible with the physical environment but are
also more or less closely ‘adjusted and fitted’ to one another, forming a
complex interweave. Any change in certain members should be accom-
panied by changes in other members with which they are linked in the
biocenosis. At that point, then, the whole biocenosis changes, in accor-
dance with the external environment, of course” [ 14, pp. 520-211.
Thus, internal changes within the biocenosis are not, in and of them-
selves, capable of wholly determining the character of a change in the
system. It is also influenced by factors that are external with respect to
it. “Any change in a species must be tied to changes not only in factors
of the external physical environment but also in other components of
the community. Only under these conditions can evolution take place,
because every organism is a part of both the living and the nonliving
environment” [ 14, p. 5201. Processes of macroevolution must be viewed
 18 RUSSIAN STUDIES IN HISTORY

in a similar context. They too must fit in with the biocenosis and the
“biocenosis-environment” system. Both microevolution and macro-
evolution form a unity with the evolution of ecosystems. “The evolu-
tion of species and other systematic groups did not take place as an
independent evolution of individual organic forms but as a part of an
elaborate community of the process of simultaneous changes in the
physical environment and biocenoses” [14, p. 5211.
Hence, evolutionaryecology made it obvious that any study of species-
formation and macmevolution must be conducted on a biocenotic foun-
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dation. In order to realize this concept in practice it was necessary to

show clearly which laws govern the evolution of communities and
ecosystems. Kashkarov illustrated how the evolution of species and
superspecies taxons depends on the evolution of the biocenosis, using
the example of the evolution of the horse. However, not many data of this
type were available in those years. In his quest for an accessible method
of studying the evolution of ecosystems Kashkarov resorted to a thought
experiment, choosing as his model a chart of the food chain in a
deciduous forest and in the tundra. Assuming that one member of the
biocenosis experienced a certain change, he traced the way in which
the entire chain and, in the long run, the entire biocenosis might be
transformed in this case, and the course of its evolution. The conclusion
he reached was this: “It is completely inconceivable that one member of
a community might evolve without others taking part” [ 14, p. 5261.
Kashkarov devoted considerable space to a discussion of the role of
isolation in the evolutionary process. He distinguished geographic from
ecological isolation and rated the latter as one of the most important
factors in species-formation. From his point of view, isolation was
always determined by ecological conditions at first. “Ecoisolates lead
to isolates that have a genetic foundation-that is, to genotypes” [9, p.
2941. Ecological isolation emerges “without the spatial separation of
populations” [ibid.]. Populations inhabiting the same area may become
isolated from one another because “they are out and about during
different seasons, they mate at different times, they keep to different
biotypes, they proliferate in different places, they encounter particular
obstacles to interbreeding with members of other populations and are
incapable of fertilizing them because of mechanical or physiological
factors” [ibid.]. As a result, the process of differentiationof a species is
accelerated, and the species separates into a number of small system-
atic units.
 FALL 1995 19

Kashkarov’s systemic approach to the study of the ecology of mod-

em forms naturally led him to problems of global ecology. He placed
the various ecological subunits, from habitat and biotype to biocenosis,
on a planetary framework. As early as 1929, in their exploration of the
question of biocenotic factors in the existence of mammals, Kashkarov
and Stanchinskii had noted: “During its lifetime an animal depends on
a number of physical factors, which combine in different ways in the
various parts of the biosphere. This results in the creation of different
habitats and biotypes” [16, p. 4571. The two became increasingly con-
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vinced that much of the data on the ecology of modem forms could
be understood “by studying the distribution of animals in its histori-
cal perspective” 116, p. 4761. Thus, they noted that various groups of
organisms (birds, for example), when adapting to identical conditions
of existence under identical ecological conditions but within the bound-
aries of isolated fauna, form “parallel ecological series o r . . . vicari-
ous groups (species, genera, families)” [16, p. 3451. In cases where
formerly separated regions of dry land entered a new conjunction
during the course of geological history, fauna would be forced to
mingle, and forms that had become adapted to identical conditions
would have to engage in a heightened struggle for existence, in which
the less well-adapted perished. Over the course of geologic time, not
only the outlines of the continents and their climatic conditions
changed but also the distribution of habitats and the composition of
biocenoses.
In practice, the principles of the global ecological approach meant
that by making a detailed study of individual fauna and comparing
them, both with one another and with the fauna of previous geological
periods, it became possible “to explore the common development of
the fauna of considerable territories and to trace their history” [16, p.
4761. Fauna characterized by endemic representatives of subclasses,
orders, and families had “long since taken an independent line of de-
velopment and were separated from other fauna” [ibid.]. Such fauna
were called kingdoms. Each of the three designated kingdoms-the
Notogaeaic, Neogaeaic, and Arctogaeaic-had developed in isolation
from ancient times. On the basis of historical considerations, the king-
doms were in tum subdivided into regions, divisions, subregions, prov-
inces, and areas [uchasfki].“Ecologically, modem conditions play no
role here” [ibid.]. In other words, the previous evolution of ecologi-
cal communities can hardly fail to exert an influence on modem
 20 RUSSIAN STUDIES IN HISTORY

ecosystems and their evolution. Consequently, when exploring the

structure of nature today it is necessary to take into account not only
the fate of individual species but also the patterns of global evolution-
ary ecology.
In affirming the biospherocentric view of the natural world, Kashka-
rov and Stanchinskii examined the role of several major taxons-fish,
amphibians, and reptiles-in biocenoses and in the general cycle of
substances in nature. In the process, they noticed that this role changed
during various stages of development of the organic world. “The role
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of any group or any species in communities and in the economy of the

entire natural world is determined by its population size or its place in
the food cycle” [ 17, p. 4451.
A comparison between the cold desert of Central Asia and the Arc-
tic enabled Kashkarov to discover an ecological parallelism between
them, one created by the similarity of several conditions of life. Having
established the composition and origin of the fauna of these regions,
and also the composition of their flora, he called them “a little
page from the postglacial period or perhaps even the glacial pe-
riod, a page from the distant past that has been preserved by excep-
tional climatic conditions-a kind of model or simulation of that
distant epoch” [18, p. 821. The natural world reacted to the onset of the
Ice Age “with profound changes in organisms and their movements in
terms of space, and with changes in behavior and instincts” [ 18, p. 831.
The biospheral approach to the ecology of deserts made it possible
to determine the place occupied by the deserts of Central Asia among
the deserts of the globe. Two types of deserts were distinguished in
Turkestan: the Middle Asian [sredneaziatskii] and the Central Asian
[rsenrral ‘noaziarskii],distinct from one another in their physical condi-
tions, their animal and plant world, and their soil cover. Preserving
common ecological features as they do, these two types of deserts
constitute “arenas in which the development of organic life proceeded
in the past and proceeds now, possibly independently” [19, p. 3061.
This, then, served to confirm the biocenological and, more broadly, the
biospheral approach to the evolutionary process. The integrated eco-
logical study of regions with many and with few changes was utilized
in this way to model the evolutionary process over considerable inter-
vals of geologic time.
Kashkarov’s ecological vision, and in particular his views on evolu-
tionary ecology, were the target of attacks as savage as they were
 FALL 1995 21

unjustified. Kashkarov was denounced for his idea that the biocenosis
is an organized, integral system, for his belief in the possibility of the
evolution of the biocenosis as a whole, and, finally, for his conviction
that the evolution of species proceeds within the confines of bioceno-
ses. He was accused of replacing “the dialectical method with the
doctrine of dynamic equilibrium,” while simultaneously it was empha-
sized that the theory of equilibrium was nourished by idealistic roots
[20, p. 81. Discussing the idea that connections exist among the com-
ponents of a biocenosis and specifically targeting Kashkarov, the cnt-
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ics wrote: “In the dynamics of individual components of a biocenosis,

these connections play a totally insignificant role in most cases, and in
theoretical terms such an assumption leads to the denial of the evolu-
tion of a species without the evolution of the biocenosis, as has hap-
pened with Kashkarov” [20, p. 71. The principal source of “errors” like
these, the critics said, was V.I. Vemadskii and his “vitalistic” postulate
that life is a dynamically steady, integral system.
Today, half a century after this criticism was voiced, not only are
ecology and Vernadskii’s theory of the biosphere being developed in
close conjunction [21], but the world scientific community is becom-
ing increasingly convinced that this is the path to take in the search for
solutions to global ecological problems and for ways to ensure the
survival of humankind [22]. It certainly deserves mention here that the
first steps toward synthesizing ecology and the theory of the biosphere
were taken in our country in the 1920s, immediately after the publica-
tion of V.I. Vemadskii’s Biosphere [Biosfera] in 1926, by V.V.
Stanchinskii [23] and V.N. Beklemishev [24].
In the half century since the death of D.N. Kashkarov, the science of
ecology has changed beyond all recognition. It has been enriched with
new approaches, new ideas and conceptions, and improved methods of
research. Nevertheless, the contributions made by ecologists of preced-
ing generations have not lost their value. A conceptual analysis of their
theoretical legacy, which incorporates a number of fruitful ideas and
generalizations, would undoubtedly be useful today. Kashkamv’s scien-
tific legacy, especially his views on evolutionary ecology, are a shining
example of this. Present-day biologists, groping their way toward a
new evolutionary synthesis, cannot fail to be interested in the accu-
mulated-but, unfortunately, as yet unrealized-experience that our
country’s outstanding ecologists gathered as they pursued their scien-
tific quest precisely along these lines.
 22 RUSSIAN STUDIES IN HISTORY

Editor’s Note

a. Further information on D.N. Kashkarov may be found in Rakhimber Um-

arbekovich Rakhimbekov, Daniil Nikolaevich Kashkarov, 1878-1941 (MOSCOW:
Nauka, 1990). and my Models of Nature: Ecology, Conservation and Cultural
Revolution in Soviet Russia (Bloomington: Indiana University Press, 1988).

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