N.K.Vereshchagin
THE MAMMALS OF THE GAUCASUS
A History of the Evolution of the Fauna
TRANSLATED FROM RUSSIAN
Published for the Smithsonian Institution, U.S.A.
and the National Science Foundation, Washington, D.C.
by the Israel Program for Scientific Translations
AKADEMIYA NAUK SSSR. ZOOLOGICHESKII INSTITUT
AKADEMIYA NAUK AZERBAIDZHANSKOI SSR
Academy of Sciences of the USSR. Zoological Institute. Academy
of Sciences of the Azerbaidzhan SSR
М.К. VERESHCHAGIN
THE MAMMALS
OF THE CAUCASUS
A History of the Evolution of the Fauna
(Mlekopitayushchie Kavkaza )(Istoriya formirovaniya fauny)
Chief Editor: E.N. Pavlovskii
Editor of this volume. A.A. Strelkov
Izdatel'stvo Akademii Nauk SSSR
Moskva-Leningrad 1959
Translated from Russian
Israel Program for Scientific Translations
Jerusalem 1967
TT66-51143
Published Pursuant to an Agreement with
THE SMITHSONIAN INSTITUTION, U.S.A.
and
THE NATIONAL SCIENCE FOUNDATION, WASHINGTON, D.C.
Copyright © 1967
Israel Program for Scientific Translations Ltd.
IPST Cat. No, 1704
Translated by Dr. A. Lerman and B. Rabinovich
Edited by M. Loeb
Printed in Jerusalem by S. Monson
Binding: Wiener Bindery Ltd. Jerusalem
Available from the
U.S. DEPARTMENT OF COMMERCE
Clearinghouse for Federal Scientific and Technical Information
Springfield, Va. 22151
1Х/9/5
TABLE OF CONTENTS
Part One. PALEONTOLOGICAL AND ARCHAEOLOGICAL
BACKGROUND OF FAUNA FORMATION
Chapter I. Development of Caucasian Landscapes and Mammalian
ELAR NTN SME TAUCIE ANC о delip sas! us. GS wit oat Mayen она
Chapter II. Development of Caucasian Landscapes and Mammalian
ЕО shaves (ONG АЕ ВЕНУ ое DR Ee О SNL oon корь ИА
Part Two. ANALYSIS OF THE ORIGIN OF CAUCASIAN
QUATERNARY MAMMALS IN RELATION TO THEIR
DISTRIBUTION, ECOLOGY, AND MORPHOGENESIS
Chapter III. Review of the Early and Contemporary Populations and
RAMGSS CH SCAMS) Па A Eis bho go ab ici oie dbo 6) ole 6 Olen
Chapter IV. Stratigraphic and Geographic Variation in Caucasian
Smareimanye MianaiialSae о Cee. Gilet neem lca ale eu ahe
Part Three. GEOGRAPHIC ZONATION AND LATEST EVOLUTIONARY
TRENDS OF MAMMALIAN FAUNA OF THE CAUCASIAN ISTHMUS
Chapter V. Analysis of Holocene Mammalian Fauna and Scheme
Oli ZONE LO) AIE Dy ET cv ON UO В rae TS ge Stra CIO
Chapter VI. Anthropogenic Changes in Mammalian Ecological
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ОА тат: 5. МОЕ СИИ, Бо Оси: lie. ok
Supplement. Part 1. Yield and Supply of Animal Pelts on the
Caucasian Isthmus from 1925 to 1955(Graphs 1-23) ........
Supplement. Part 2. Maps of Animal Distribution on the Caucasus
арены ее кое аа
Index of Common Names of Mammals........... О о
Index of Latin Names of Матта1 ..... В А
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English page
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PREFACE
N.K. Vereshchagin's book is somewhat unusual: although it is not
a comprehensive treatise of the past and present evolution of the fauna of
this interesting and complex area, he has succeeded in producing the first
generalized and complete picture of the development of terrestrial fauna in
the Caucasus from Neogene to Recent times against a background of the
geological history and landscape shifts.
In his work the author draws on the entire body of biological science,
from paleontological data on individual stages in the phylogeny of Quaternary
mammals to the voluminous data of modern zoogeographical and ecological
conditions of contemporary forms.
The author believes that the eastern Mediterranean (including the
Caucasus) was characterized in the Neogene by a unique process in the
mammalian evolution — faunal complexes replacing one another, as
commonly occurred throughout the Recent in the Holarctic region.
The Quaternary mammalian faunas of the Caucasus have their roots deep
in the Pliocene. The Caucasian faunas proper of the mountain forest
community evolved locally since at least Upper Miocene time. However,
in the Quaternary the evolution of mammals in the Caucasus took place
primarily through species invasions and extinctions.
The evolution of new species on the Caucasian Isthmus was not as
Significant as other evolutionary processes in fauna formation.
Veresnchagin's statements on the absence of saltations in the
morphological evolution of the Quaternary mammals will require additional
study before they are confirmed.
Emphasis is placed on the destructive activity of humans which caused
the extinction of many game mammals from the Upper Paleolithic to the
Recent. Many examples are given of the ecological changes and
readjustments of the surviving mammals in various zones.
This work on the many questions relating to the development of fauna
will undoubtedly inspire other intensive paleontological and zoological
studies, not only on the Caucasus, but also on other parts of Russia. As an
initiator, Vereshchagin has made a notable contribution to the methodology
of his subject.
The book also presents important data on the commercial value of
autochthonous and introduced species, as well as information of concern
to conservationists.
The history of the development of mammalian fauna in the Caucasus will
undoubtedly be of interest to a wide audience of botanists, geologists,
archaeologists and geographers, as well as to zoologists.
E.N. Pavlovskii, Academician,
Lieutenant General, Medical Corps.
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INTRODUCTION
A study of the distribution of organisms in time and space in the course
of their evolution presents problems at once interesting and difficult. Their
study is necessary for the reconstruction of faunal history and the
advancement of faunalogical investigations.
The study of the development of fauna on the Caucasian Isthmus is
complicated by the fact that throughout the Cenozoic, this area was the
scene of marine transgressions and regressions, mountain-building
processes and glaciations. As a result of these processes, the climate and
landscape changed within short distances.
In the physiographic sense, the Caucasus is large and varied. Its geologic
structures, its flora and fauna, and the history of its peoples have been
studied by outstanding scientists for almost 250 years.
Studies of the mammalian fauna of the Caucasus have been primarily
concerned with the identification and geographic distribution of species.
Few workers were concerned with the ecology, and even fewer with the
history of the fauna. The main works are those of K.A. Satunin,
N. Ya. Dinnik, N.A. Smirnov, M.V. Shidlovskii, Z.S. Rodionov,
В. А. oviridenko,, 5. 1. Орех, ‚т. В. Beme,) 5.5, Murov;,(V)G..Geptner,
ВТ. Ако1корию, А. А. Nasimovich, 5. №. Ша! Т.М. Яваскох ааЯ
Р.Р. Gambaryan.
The vast collection of data by Russian zoologists on Caucasian animals
can only be partly reflected in the bibliography of this book.
Much paleontological work on the fossil mammals of the Caucasus
has been done by V.V. Bogachev, A.A. Borisyak, V.I. Gromov,
Е.Г. Belyaeva, У.Т. Gromova, В.О. Dzhafarov, L.K. Gabuniya,
N.O. Burchak-Abramovich, myself and others. The osteological collections
of archaeologists (S.N. Zamyatnin, G.K. Nioradze, A.P. Kruglov,
Е.Г. Krupnov and others) were of particular value.
These studies and collections facilitated my investigations of the origin
and history of the development of Caucasian mammals.
The history of the mammalian fauna of the Caucasus is intimately related
to the evolution of landscapes, vital forms and ranges of species distribution.
The marine transgressions and regressions during the Tertiary in the
eastern part of the Mediterranean geosyncline (the Recent Black Sea,
Caspian Sea and the Caucasus) and the evolution of the molluscan fauna of
this area have been studied in great detail by geologists: Academicians
Andrusov (1888) and Arkhangel'skii (1934), Arkhangel'skii and Strakhov
(1938), Zhizhchenko, Kolesnikov, Eberzin (1940) and others, mainly in
the course of petroleum explorations. The Neogene, inparticular, has been
thoroughly studied. A detailed zoogeographic summary оп the faunas of the
Caspian and Black seas was published by Sovinskii (1904).
Geobotanists and paleobotanists have also published a number of histories
of the land flora of the Caspian lands, particularly of the Caucasus
(N.I. Kuznetsov, 1909; Palibin, 1936; Grossgeim, 1936, 1948; Maleev,
1941; and others).
Studies to date on the history of Caucasian land fauna have been
quite generalized and were based mostly on fossil material. These studies
include Academician Menzbir's (1934) history of the fauna of the European
U.S.S.R., Serebrovskii's (1935) history of the fauna of the U.S.S.R.,
Bogachev's (1938) review of the Tertiary of the Caucasus, and Borisyak
and Belyaeva's (1948) review of the Tertiary mammalian fossil localities
in the Caucasus. The history of the Quaternary fauna has been treated by
Gromov (1939, 1948), Gromova (1948), Burchak-Abramovich (1951c) and
Pidoplichko (1951, 1954).
The history of the land mammals of the Caucasus has also been treated
from the point of view of zoogeographical studies of the Recent by Satunin
(1896, 1901а, b, 1904, 1909, 1913), Dinnik (1911), and Shidlovskii (1940a,
1941b, 1945, 1947), and more recently by B. A. Kuznetsov (1949, 1950).
There are fewer studies on the evolution of other classes of terrestrial
vertebrates: birds (Puzanov, 1938b) and reptiles and amphibians (Nikol'skii,
1913; Sobolevskii, 1929; Lyaister, 1931; Chernov, 1939).
The history of terrestrial invertebrates on the Caucasus has been
discussed in the studies of scorpions by Byalynitskii-Birulya (1917)*; of
dragonflies by Bartenev (1933, 1934 — in which he also discussed mammals,
reptiles and amphibians); of Coleoptera by Semenov-Tyan-Shanskii (1936)
and Bogachev (1947); and of Orthoptera by Uvarov (1921).
All of these studies have one feature in common: the history of the fauna
is treated either from a purely paleontological or from a purely
zoogeographical point of view, without synthesis or consideration of the
ecological and morphophysiological data.
Obviously, this approachcannot give a true picture of a process as
complex as the development of fauna at any given stage of geologic time.
It is more correct to view the process of the development of fauna as
a combination of three interrelated processes:
1. Evolution of the environment and of those life conditions of individual
species which were controlled by secular changes in climate, in elevation
and subsidence of the earth's crust, andofthe biocenosis proper. In more
recent times the human factor also assumes great importance.
2. Morphological evolution of individual species, which is affected by
both external environment and internal developmental patterns.
3. Ecological evolution of individual species and of entire biocenoses
which is caused by continuous evolution of the conditions of life, and the
further evolution of the biocenoses, in turn, through extinctions,
transformations, invasions and migrations of species.
Data on these three processes of faunal evolution are not equally
available to paleontologists, zoogeographers and ecologists. A synthesis
of the data and conclusions of each discipline is indispensable.
The Caucasus has had a complicated geological, pedological and floral
history, and a long pattern of changing human cultures. The influence of
* [Arthrogastran Arachnids of Caucasia: Scorpions, Translated into English by IPST in 1964, OTS No, 64-1114.]
man оп the land and the fauna became progressively more pronounced with
time. Therefore, in order to understand the basic features of the evolution
of Caucasian fauna, it is necessary to draw on the data of geology,
geomorphology, geobotany, archaeology, paleontology, systematics, ecology
and zoogeography.
The complexity of the problem required a limitation on our studies and
an organization into three interrelated subdivisions:
1. Paleonio!ogical and archaeological evidence of the history of the
development of the fauna.
2. Analysis of the origins and distribution of the Quaternary mammals
with reference to their ecology and morphogenesis.
3. The mammalian geography of the Caucasian Isthmus and the most
recent manifestations of local faunal evolution.
In each of the subdivisions, progress of the work depended upon the
material and technical facilities available.
Materials, routes and methods
The main source of mammalian material of the Tertiary and Pleistocene
is in the paleontological, geological and archaeological collections of:
Museum of Natural History of the Academy of Sciences of the Azerbaidzhan
S.S.R.; Zoological (ZIN) and Paleontological (PIN) Institutes of the Academy
of Sciences of the U.S.S.R.; All-Union Geological Institute, Institute of
Geology of the Academy of Sciences of the Georgian $.S.R.; Institute of
Geology of the Academy of Sciences of the Armenian 5.5.В.; and the city
museums of Pyatigorsk, Stavropol, Krasnodar, Temryuk, Novocherkassk,
Rostov and Astrakhan. Our own collections and observations on Tertiary
mammals were taken from the Miocene site near Belomechetskaya in
central Ciscaucasia, from the Pliocene site at the Kosyakin quarry near
Stavropol and from the Upper Pliocene and Lower Quaternary sites along
the Psekups and on the Taman Peninsula (Figure 1). Collections of
Pleistocene mammals and plants were taken from Pleistocene river sands
in the Kuban and the Kuma river valleys, near Krapotkin and Georgievsk
in Ciscaucasia, and from the bituminous formations of the Apsheron
Peninsula in Transcaucasia (Figure 2).
Small collections of bones and flint tools and additional observations were
made at old excavation sites in paleolithic caves on the Black Sea coast
and inthe Rion River basin(Akhshtyrskaya, Sakazhia and Gvardzhilas caves)
and in the recently excavated Kudaro cave.
More than 55,000 Pleistocene bone fragments and 1,500 Tertiary bone
fragments were examined.
In order to trace the distribution of game animals and small rodents
in the Holocene, the following types of fossil material from a variety of
sites were studied:
1. Food and industrial wastes of man from strata containing remains
of campsites and settlements from the Mesolithic to the last centuries of
the Recent (Sarkel, the ancient town of Semibratnoe, Elizavetovskoe,
Cepi, Phanagoria, Taman, Gelendzhik, Anaklia, Kalakent, Mingechaur,
Baku, and others).
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domestic animals from ritual burial grounds (Nal'chik, Samtavro, Trialeti,
Sevan and others).
3. Skulls of artiodactyls from mountain caves and shelters where they
had accumulated over the centuries as the residue of rites and feasts
dedicated to the hunting god(a number of ravines in Ossetia).
4. Bones of rodents and insectivores deposited by owls during the last
centuries in caves, under ledges and in the small cavities of rocks (the
foothills and mountain regions of Cis- and Transcaucasia).
5. Bones and horns of drowned animals and kitchen middens of tribes
which lived in pile dwellings along the shores of Lake Sevan. (The lake is
at present receding rapidly. )
6. Isolated occurrences of bones in Holocene loams.
In all, nearly 52,000 identifiable bones and bone fragments from
approximately 70 Holocene sites (Figure 3) were studied.
Penetration of species in postglacial times and their role in the evolution
of faunal complexes were the primary considerations in our selection of
smaller areas of Holocene localities such as the Pleistocene localities of
the Apsheron Peninsula, Stavropol Plateau and Pyatigor'e area
(Vereshchagin, 1949c, 1953a). We particularly searched out food rests of
eagle owls.
Mapping of the fossil localities and searches in the extensive literature
produced a general picture of the distribution of bone-bearing deposits of
the Tertiary and Quaternary.
The elevation of the Caucasus in the Cenozoic resulted in an accretion
of very thick strata of gravels, sands and silts in the piedmont plains. Part
of the terrigenous material was reworked by waves in the surf zone and
part was deposited in the quiet water of bays on the margins of sea basins.
Accumulation and burial of skeletons occurred mainly along the deltas
and coastal bays, where they were carried by streams. Cenozoic bone-
bearing sandy-gravelly lenses are usually found in erosional channels cut
through older beds; this type of burial is common in the valleys of
Ciscaucasia, Stavropol and in the broad intermontane valleys of
Transcaucasia. The bone-bearing formations are exposed in the processes
of erosion and quarrying.
Caves containing bones of Tertiary age have not yet been found in the
Caucasus. This is probably accounted for by the changes in the relief and
river network which destroyed the older karst.
Archaeologists have searched for Paleolithic localities inthe regions of
developed Quaternary karst. The Paleolithic collections usually contain
"fauna", i.e., bones left in the caves by prehistoric hunters and by
predatory animals and birds. The latest, post-Paleolithic (post - Pleistocene)
bones occur mainly in surface loams, diluvial gulleys, alluvial deposits
in the first terraces of rivers, in caves and under rocky ledges.
The positioning of most Holocene bone-bearing burials accessible to
excavation and investigation was determined by early hunters and predatory
birds. However, evidence of mass extinction of wild animals and
preservation of their remains even in Recent times can be found in the
Caucasus (Vereshchagin, 1951b; Figure 4).
It is clear that the paleontological record of the origin of Caucasian
mountain fauna (which is the background of this work) is not complete.
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The reasons for this are taphonomical, i.e., the lack of accessible areas of
sediment accumulations inthe highlands and the absence of caves containing
Pliocene fossils. (See Efremov: ''Taphonomy'' (Tafonomiya) 1950. *)
A critical evaluation of the available paleontological material requires a
complete account of burials, their peculiarities and the ways by which bone
remnants and matter were preserved in them. Therefore, a section dealing
with the transport pathways and theburial of fossils has been added to
each of the chapters devoted to the regional stratigraphy of the localities.
A correct evaluation of the modes of preservation leads to important
conclusions regarding the ancient landscapes and the conditions of life and
of death of organisms. However, this factor was often ignored by
paleontologists and, as a result, they were led to erroneous interpretations,
particularly in cases where fossils of varying ages occur at the same
locality.
In most instances, the paleontologist deals with a collection of fossil
remains which reflects only the conditions peculiar to a particular locality,
rather than the ecological associations of the entire ancient region.
The degree of "universality'' of a burial is related to the number of
preservable species of animals and plants which inhabit the area and to
the proximal degree to which the distribution of species and individual
specimens in the dead assemblage resembles that of the living community.
Three sets of conditions determine the degree of universality of a fossil
assemblage: 1) the circumstances ofthe animal's death; 2) the manner in which
the remains accumulated in sediment; and 3) their ''behavior'' and
preservation within the sediment.
Clearly, any possibility of reconstructing ancient biocenoses and
landscapes from fossil material is determined by the sets of conditions
given above.
The Binagady burial on the Apsheron Peninsula (Chapter II) is an example
of a highly universal fossiliferous site. There a Pleistocene assemblage of
plants, birds, insects, jerboas and rhinoceroses was preserved in thick
layers of asphalt; this mode of preservation permits a highly reliable
restoration of the ancient landscape.
The situation is quite different when fossils are preserved in sediments of
diluvial, river, lacustrine or marine origin, ог at Paleolithic sites or incaves.
Accumulation of bone material in river sediment results from river
erosion of bone-bearing beds (Vereshchagin, 1953c), as well as from the
occurrence of animal deaths in the mainstream and оп the floodplain. Once
in the mainstream, the bones are subject to mechanical reworking like
gravel and pebbles and they are sorted and distributed according to specific
gravity and size.
Thus, a complex of animals which died in a river valley can be seen
accurately only ifthe bones are collected from different facies, i.e., gravels,
sands, silts.
Available collections by earlier investigators were not, as arule, made
in this way and consequently reflect only the last stage of sorting, i.e.,
excavation.
Distribution of bones in different facies also occurs in lakes and marine
bays to which they are carried by rivers. Burial in deltaic sediments is
more common, since floating bodies of animals can be carried long
* [The term "taphonomy," meaning "the study of the formation of burials of fossils, plants and animals, "
was introduced by Efremov in this work, ]
14
distances by currents to sink later into the silty and shell-covered bottoms
of quiet bays.
Accumulation of bones in the Paleolithic and post-Paleolithic strata of
camp sites, settlements and burial grounds was primarily dependent upon
the particular hunting traditions and customs of local tribes. (Deposition
by flowing water is rarely encountered in caves. ) Remains of men's food
was often found in caves mixed with remains of the food of predatory animals
(bears, wolves, hyenas, panthers) and predatory birds (eagle owls and little
owls). Clearly, grosserrors are possible if, indrawing conclusions, human
hunting customs and animal behavior patterns are ignored, аз was done in those
investigations which are limitedto a count of domestic and wild animals instrata
of varying ages and cultures. For example, it is known that contemporary
Caucasian hunters often carry 30-40 kg of wild goat, gazelle or swine meat
over distances of 20-25 km on mountain trails. Caucasian goats can climb
1,500-2,000 m in a day. We have observed eagle owls in the cave region of
Imeretia flying frequently from caves in the foothills to seek prey on
the subalpine meadows and attaining heights of more than 1,000 m in their
ascents.
Thus, reconstructions of the paleolandscape and of changes in the
position of the snow line based on occurrences in caves of the remains
of highland animals (goat and Prometheomys ) can be made only if all
the processes involved in the accumulation of bones are considered.
Similarly, conclusions on the accumulation of index gama fauna of one
area cannot be based on a comparison of the material from an unlike area,
e.g., material from the Greek towns of the Taman Peninsula (Phanagoria,
Cepi) compared with the kitchen middens of a small hunting and fishing
village of Roman time found at the ancient site of Semibratnoe in the mouth
of the Kuban River. In this case the size of the settlement and its
topographic location would be much more important considerations than the
occurrence and disappearance of game animals in the surrounding area.
Quantitative data on the composition and size of burials in the Caucasus
are given here for the first time whenever adequate material was available.
These data contribute to an understanding of the development of tribal
culture and economy and of the nature of the landscape. To take one
example: unverified ancient writings have led some geologists to believe
that dry valleys existed in Colchis in the first millennium B.C. This theory
is disproved by the absence of horse remains and the abundance of boar
remains in the Colchis burials, which indicate that the marshlands of the
Eneolithic and Bronze Ages were very Similar to those of approximately
50 years ago.
The species composition and distribution of wild animals were also
inferred from the drawings and sculptures of the Bronze and Iron ages.
Particularly interesting are the bronze, silver and gold objects with animal
representations collected from the burials of the Armenian Highland, the
Kura River valley and the Trans-Kuban sloping plain (Tsalka, Samtavro,
Maikop, Kellermes, etc.). If one takes into account the probable origins of
these objects, the aesthetic criteria of their consumers (Urartus,
Scythians, Kobanians) and the ecology of the depicted, the realistic art of
these ancient craftsmencan be of great aid in studying the distribution and
gradual extinction of some of some of the larger animals.
Quaternary
Neolithic settlements
Holocene
(Bronze and Iron ages)
V WA Ze
Pleistocene
Asphalt pools
Tertiary
Sea bays and river deltas
FIGURE 4, Predominant circumstances of animal death and types of fossil burials in the Cenozoic of the
Caucasian Isthmus
15
Although it is possible to find fossils of large animals of the present
‘period, their number has decreased to an extent that makes this a negligible
probability. Consequently, the zoologist who is called upon by archaeology
to furnish morphological and zoogeographic data for the identification of
depicted animals will attach great importance to the so-called ''animal
style'' in the art of the first millennium B.C.
We have reviewed the published data of Tolstoi and Kondakov (1889),
Uvarova (1900), Кайт (1940) and others on precious stones in the ''animal
style'' from the Caucasian burials of the Kobanian, Khodzhalian, Scytho-
Sarmatian and other cultures, as well as the collections of the Hermitage
(Leningrad), the Georgian Museum (Tiflis) and the Historical Museum
(Moscow). Our conclusions, as they relate to species and ranges, are
discussed in Chapter Ш. The ‘animal style" of these Caucasian antiquities
deserves a special zoological-ethnographical study in itself.
In the study of the Cenozoic faunas of the Caucasus we used the
geochronological subdivision of the Neogene of the eastern Mediterranean
(Black and Caspian seas, Caucasus) as given by Academicians Andrusov
(1918) and Pavlov (1925), with later additions by Kovalevskii (1933, 1936),
Zhizhchenko, Kolesnikov and Eberzin (1940).
We placed the Pliocene-Pleistocene boundary (in agreement with Pavlov,
1936, and Vardanyants, 1948) at the end of the Apsheron stage. The
boundary probably corresponds to the last stage of existence of the Taman-
Psekups fauna in Ciscaucasia, which was originally discovered by Gromov
(1948, 1950) and is described in Chapter II of this book.
The Quaternary of the Caucasus is subdivided into the Pleistocene and
Holocene. Everywhere on the Caucasus the development of faunal complexes
during the Pleistocene extended over a fairly long period, fromthe Upper
Apsheron to the end of the last glaciation on the Caucasus and the onset of
the xerothermic period which marks the beginning of the Holocene. The
Pleistocene is subdivided into the Lower, Middle and Upper Pleistocene,
which correspond to the stratigraphic stages of the Caspian region: the
Baku, Khazar and Khvalynsk, containing mammalian fauna known from the
entire Russian plain. The subdivision of the Quaternary in the foothills of
the Caucasian Isthmus and the correlation of the river terraces with cave
deposits and marine terraces were established following the studies of
Mirchink (1937b), Gromov (1948), Vardanyants (1948) and Nikolaev (1953).
The archaeological chronology oftne Caucasus is similar to the European
(West Mediterranean) schematic chronology developed by Obermaier (1913),
Osborn (1924) and Penk (1939). For date determinations of the Caucasian Paleo-
lithic as guide lines to grouping mammalian ecological complexes, we used the
papers of Zamyatnin (1950b, 1957) andother archaeologists. Geological dating
was a primary concern in our studies of bone material.
The problem of determining the relative and absolute age of fossil bones
is becoming increasingly significant for geology, paleontology and
archaeology. Fossilage is determined by: 1) physicochemical composition
(the methods of calcination andradiocarbon dating); 2) taxonomic characters
(generic, specific, subspecific); 3) inclusion in one faunal complex or
another (faunal assemblages); 4) artifacts occurring with the fossils (stone
tools, pottery, etc.); 5) evidence of associated invertebrate fauna and
indirect geomorphological data (for example, the age of marine and river
terraces); 6) type of preservation of the bone material.
16
Each of the methods mentioned has disadvantages. The physicochemical
methods give comparable results only when all the bones studied are less
than 500,000 years old and have been preserved under the same conditions.
The factors which affect the data obtained by this method are the composition
of the sediments and their radioactivity, the water-salt relationship in the
soil at the time of burial, etc.
However, the method of calcination and comparison cf the coefficients
of the organic residue used by Pidoplichko (1952) can be used in absolute
and relative dating provided that separate chronological scales are р
constructed for each physicogeographic region and for each type of locality.
Radium-, uranium- and thorium-isotope dating methods (Cherdyntsev,
1955) yield inconsistent results.
Determination of geologic age by specific and faunal characters is
possible only if the evolution of morphological characteristics is known in
detail for the given genus or species. The taxonomic method can only be
used with extremely well-preserved material, particularly in the case of
skull specimens.
Stone tools, pottery and other artifacts are reliable indicators of the
relative age of fossiliferous strata only in those geographically defined
areas where the chronology of the human cultures is known. By the latest
consensus, the development of the Paleolithic cultures in Western Europe
did not coincide in time with the Paleolithic in Eastern Europe. The
occurrence of the bones in situ is another prerequisite for successful
application of this method.
Determination of the age of the geologic formations is of little use in
determining the age of bones in Quaternary sediments which were often
redeposited. The problem of dating is usually reversed since the Quaternary
beds are dated by the vertebrates found in them.
This method uses the simplest organoleptic analysis to determine the
mode of preservation of bone material. But it requires considerable
experience for an unbiased judgment and only yields indexes of relative age.
It was necessary to review the entire body of Quaternary paleontological
material taken from the Caucasus, since the indiscriminate application of
morphological and morphometric criteria by earlier investigators had only
resulted in confusion and redundancy in lists of species.
With careful consideration of the mode and conditions of preservation,
the age of the bone material can be placed within the major subdivisions
of the Quaternary. The loss of organic matter (fossilization) and the
secondary permineralization of the bones must also be taken into account
by observing the new saturation of the bone material by salts, the
color of fresh fracture, the degree and depth of colorization and the smell
of fresh bone when scraped with a knife. A table is given for the
determination of the relative age of Quaternary bones (Vereshchagin and
Gromoy, 1953а).
The geographic distribution of Recent species was studied, with
interruptions, from 1935 to 1941 in Azerbaidzhan and Armenia, and from
1945 to 1952 in Azerbaidzhan, Georgia, North Ossetia, Kabarda, Svanetia
and Dagestan. Our collecting routes are shown on the map (Figure 5).
Studies were made of the collections of ZIN, the Zoological Museum
of Moscow University, The Georgian Museum and Institute of Zoology
of the Georgian Academy of Sciences (collections to 1944), the
Institute of Zoology of the Armenian Academy of Sciences (collections
to 1943), the Institute of Zoology of the Azerbaidzhan Academy
14
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15
(collections to 1948) and the Stavropol and Baku anti-plague stations
(collections to 1945). A total of 13,500 excellent specimens of skins, stuffed
animals, skulls and specimens preserved in alcohol (taken from over
3,300 locations) comprised the material used.
Conclusions on the most recent changes in the density of 23 species of
fur -bearing animals were based on the data collected over 25 years (1925-
1950) by the governmental fur stations in Ciscaucasia and Transcaucasia.
17 The data of 257 pelt-receiving centers for the period 1945-1948 were used
in the final refinement of the areas of distribution of species, as Shown in
Figure 5. The statistics on the preparation of furs are shown separately on
the graphs for Ciscaucasia and Transcaucasia, and for the republics and
regions (except the Astrakhan Region, for which no data were available).
The occurrences of fossils and the recent yield of small fur-bearing
animals are entered on a specially prepared map drawn to a scale of
1:5,000,000. The population density of each species, as estimated from the
mean annual number of furs received by the pelt-receiving centers, are
indicated by small dots on the map. The total number of pelts exceeds
3,665,000. Range boundaries have been drawn only for stenotopic species
(i.e., those with a narrow geographic range). The occurrences of widely
distributed animals, such as boar, gazelle, deer, antelope, saiga, goat and
otter, are given according to our own observations.
Data from the literature, not confirmed by observation, are entered
with a query.
The history of the development and the present state of distribution are
given only for the stenotopic species and for those which are well
represented in the paleontological record.
The history of animal distribution in the Caucasus is closely connected
with the climatic zones of the Quaternary. A relationship between vegetation
and species distribution is also valid for this period during which there
was a Successive displacement of xerophytic desert vegetation, first by
a steppe landscape, then by a mesophytic, andlastlyby a forest landscape
followed by a reversal of the entire process of vegetation displacement.
The relationship, however, is not as clear in the older periods. It is certain
that conclusions on distribution areas are easier to reach and are more
reliable, asthe species association with a given type of vegetation and
landscape is longer and its ecologic niche is more limited.
Gaps in distribution and the influence of ecological barriers are only
considered as they relate to types of vegetation and to barriers which were
geophysical and climatic in nature. Ecological barriers which are
determined by physiological reaction norms and morphological adaptations
of the species require additional study. The validity of our theoretical
assumptions was confirmed by the finding of the indicator species
associated with a given type of vegetation and landscape in every case.
The morphology was studied mainly from the Quaternary fossil materialin
relationship {о {$ stratigraphic distribution andthe rate of evolution ofthe spe-
cies. The morphological studies were primarily of carnivores (Vereshchagin,
1951b) and of some rodents and hoofed species (Chapter IV). The geographic
variability of some Recent carnivores, ungulates and rodents was also
18 investigated in order to understand their origins and geologic age.
16
1)
The taxonomic classification of fragmentary bones of Quaternary
mammals proved to be a problem. Some mammals, known from the Middle
Pleistocene to the Recent, show gradually evolving successive forms with
very few morphological differences. Other lineages of mammals are
represented by a succession of more or less distinguishable forms which
can be discussed as subspecies of the Recent orfossilspecies, or as species.
As arule, paleontologists either classify the Upper Quaternary forms
as subspecies of existing species, or they identify the fossil with the Recent
species. This identification of a fossil form as a subspecies of a Recent
species is unsound, as the criteria for subspecies distinction are quite often
only conventional and subjective.
Although we are out of agreement with this practice of classification,
we retained in Tables 62 and 103 the names of the subspecies found in the
literature. Those subspecies whose stratigraphic records are inadequate
are entered with the qualifications conformis (similar) and affinis
(related). Unfortunately, the Rules of Zoological Nomenclature (1932)
do not cover this problem.
Wherever possible, we applied ecological data of the Recent to the past,
and the study of the ''universal'' death assemblage at Binagady proved the
effectiveness of this approach.
Although a certain constancy of predominant features is recognizable
in Quaternary organisms, it is probable that there was a fairly rapid and
extensive evolution of the physiology. This brought about modifications in
the mode of life without changing the former morphological features.
The zoogeographical analysis of the entire fauna of the Holocene mammals
of the Caucasus is the culmination of studies of the history of faunal
complexes. The analysis is based on genetic and stratigraphic principles.
Rather than adhering to geomorphological and phytogeographical units, we
have emphasized the evolutionary aspects of areas of species distribution
and of faunal complexes.
The problem of the geographic origins of species is discussed in
Chapter III along the lines of criteria proposed by Arldt (1919).
In any discussion of the future evolution of ranges, the chief consideration
for any geological stage is undoubtedly the human factor. With this in
mind, we studied the numbers and behavior of animals in their original
biotopes and in those influenced by man, selecting various geographic zones
mainly in the eastern Caucasus.
These investigations add to our understanding of recent evolutionary
development or degradation and direction of the fauna.
Quantitative studies of animal populations in nature preserves were
carried out during the expeditions (1938-1948) of the Azerbaidzhan anti-
plague station and the Institute of Zoology of the Academy of Sciences of
the AzerbaidzhanS.S.R. Most of the studies employed commonly known
methods, although in some cases special new methods nad to be devised.
Studies of recent changes in virgin steppelands were done mainly in
Azerbaidzhan and should be carried out in the future in Ciscaucasia where
shelterbelts have been planted.
Introduction by man of new species into the existing fauna is not
discussed in detail in this book. Acclimatization of new species and the
biotechnical procedures of game farming, which have been studied by many
scientists, are separate problems. Nevertheless, some examples are given
20
in this book of the rates and patterns of distribution of some introduced
carnivores and rodents. *
The effects of man's alterations of the landscape were clearly observable
in the most recent changes in ecological assemblages; in fact, they may
completely overtake the natural processes of change. For example, the
development of stunted thorny shrubs in the place of forest in the foothills
of eastern Ciscaucasia is a result not only of the Recent climatic trend, but
also of deforestation and cattle grazing. Another example is the artificial
drainage and forestation of the central parts of the Colchis Plain, which
completely suppresses the development of water-logged soils and hydrophilic
vegetation. In both cases the original assemblages of large and small mammals
were completely destroyed.
The following ecological and zoogeographical terms are used throughout
the book:
Ecological assemblage of mammals — a group of species which inhabit
one biotope. The morphological and physiological features of the species
are the result of evolution within the framework of the existing ecological
conditions.
Faunal complex — a number of ecological assemblages occurring within
one homogeneous geographic zone.
For example, the faunal complex of the arid eastern Transcaucasian
plains from the Pleistocene to the Recent includes ecological assemblages
types. This complex has evolved since the Pliocene under dry and
moderately warm climatic conditions.
The term 'fauna' is used in the book to designate a number of faunal
complexes occurring in a multizonal territory. The word 'fauna' is
commonly used by geologists, paleontologists and archaeologists to designate
any collection of fossils. We do not use the word in this sense: where it was
necessary to employ it, it appears in quotation marks ("'fauna''). It is a
mistake to identify a selected collection of fossil remains of animals,
possibly of different ages, with the fauna, or the faunal complex, ofa given
region or country.
Our terms are quite applicable to paleogeography, i.e., to the older
stratigraphic stages. A faunal complex can exist throughout an entire
geological epoch, during which its composition will gradually evolve. The
extent to which it is possible to reconstruct fossil ecological assemblages
and faunal complexes depends upon the degree of completeness of the
geologicalrecord. It is for this reason that index species which occur in large
populations at certain stratigraphic horizons are indispensable to a
reconstruction.
From the occurrences of such index species of the Pleistocene steppes
of Eurasia as mammoth, horse, bison, saiga and other species, it is
possible to infer the occurrence also of suslik, marmot, corsac fox,
Siberian polecat, steppe skunk and related species. Similar inferences
can be made from the occurrences of mountainous index species.
In the process of preparation of this book, it was necessary to use
extensive paleontological, zoological, botanical, geological and
archaeological literature. Ofthe more than 3,200 sources consulted, only
the most frequently quoted are given in the bibliography.
* The author participated in the introduction of nutria, common raccoon and mink into the Caucasian fauna
and served as a consultant in a number of other experiments.
** [Tugai — a bottomland complex with forests, bushes and meadows in river valleys of Central Asia. ]
18
21
The writings of ancient Greek, Roman, Arab and Armenian naturalists,
historians and geographers in translation, as well as the reviews of Gan
(1884-1890), Latyshev (1893-1904, 1947, 1948), and Karaulov (1901)
were used as source material.
Of the voluminous literature on species systematics only the titles most
essential to the Caucasian fauna are given.
The Caucasus (Caucasian Isthmus) is considered to lie within the
following boundaries: the Kuma and Manych rivers to the north, and the
international boundary of the Trancaucasian republics to the south.
Of the many questions and problems which arose during the course of
the field work and preparation of the book, some of the most important
involved the stratigraphic correlations of paleolithic localities in mountainous
country wherein several climatic, floral and faunal provinces are joined.
The present work could have been accomplished only with the support
of the Directors of the Zoological Institute of the Academy of Sciences of
the U.S.S.R., and the Presidium of the Azerbaidzhan Academy of Sciences.
The author is indebted to B.S. Vinogradov and A.A. Strelkov for valuable
editorial comments and advice.
The line drawings were done by V.N. Lyakhov, the drawings of animals
by E. Ya. Zakharov and Prof. К.К. Flerov. All the photographs were taken
by the author.
The interest and cooperation of officialinstitutions and individuals, both
specialists and laymen, was invaluable to the field work in Azerbaidzhan,
Armenia, Georgia, Dagestan, North Ossetia, Stavropol and Krasnodar.
I thank all those who, in one way or another, contributed to the
completion of my work.
The first and more detailed manuscript was completed in 1954; in the
process of preparation for this publication, many sections were shortened
and new material was included.
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25 Part One
PALEONTOLOGICAL AND ARCHAEOLOGICAL
BACKGROUND OF FAUNA FORMATION
Chapter I
DEVELOPMENT OF CAUCASIAN LANDSCAPES AND
MAMMALIAN FAUNA IN THE TERTIARY
ТВ Ви S@ ONE - BEARING Si@i@ Agi aa S
Abundant remains of Carboniferous plants in the slates and sandstones
along the northern slopes of the Main Range indicate that land existed in
the Caucasus as far back as the Paleozoic.
Individual islands appeared after the broad transgressions of the
Mesozoic, and later merged gradually into one landmass.
FIGURE 6, Land formations (cross-hatched ) in Eastern Europe
and Southwest Asia (from Arkhangel'skii and Strakhoy, 1938)
Dot indicates Upper Oligocene locality at Benara
The Caucasian islands were covered by ferns, ginkgos and cycads in
the Jurassic, but the paleobotanical record only commences with the
Cretaceous when plane, poplar, dryandra, myrica and giant conifers
(araucarias, sequoias and others) appeared.
Iguanodons which occur in travertine encrustations in the Satapliya
cave near Kutaisi are the only fossils representative of the terrestrial
vertebrates of the islands (Kandelaki and Dzontsenidze, 1937; Gabuniya,
1956b).
According to Pavlov (1936), the area between southwest Asia and the
Baltic was dry land in the Paleocene. The Caucasus was built of gently
folded Jurassic and Cretaceous strata.
2
At a later stage, the transgression of Tethys covered southern
Europe, the Ukraine, the Crimea and part of the Caucasus, and Tethys
became connected with the Asian sea extending to the southeast.
Thus, the formation of land in the Caucasus, which determined the
evolution of the Quaternary faunas, took place entirely during the Cenozoic.
In the Eocene the northern coast of Asia Minor, Surami, Kazbek and
Elburz were sites of intensive vulcanism, which probably produced the
high relief of the island lands of the Caucasus (Figure 6). There are no
data on the Eocene flora.
Small whales of the genus Zeuglodon (Archaeoceti) lived in the sea
surrounding the island of Caucasus. Their fossilized remains occur in the
Koun beds of the Apsheron Peninsula and in the area of the Sumgait rivulet.
The first occurrences of terrestrial vertebrates in the Caucasus are
in the Oligocene.
OLIGOCENE
The tropical character of the Caucasian landscape persisted through
the Oligocene. The tuffaceous sandstones of Lower Oligocene age in
Mount Darry-Dag and the Araks valley contain ferns (Blechnum brauni,
Pteris oeningensis), palms (Sabal hearingiana), conifers
(Podocarpus), Lauraceae (Cinnamomum), myrica, zelkova, and
small grasses (Panicum miocenica and other forms). Palibin (1936)
correlated the Darry-Dag Oligocene flora with the Lower Oligocene floras
of the Balkans and central Europe. He thought that the Darry-Dag is
indicative of humid tropical forests covering volcanic islands.
No bones have as yet been found in the Lower Oligocene beds of the
Caucasus. However, it is possible that the island of Caucasus and the
adjacent land masses to the south were inhabited by the same fauna of
primitive carnivores and hoofed mammals which are well-known from the
Eocene and Oligocene deposits of Western Europe.
Bogachev (1938d) mentions a footprint of а creodont about the size of a
large dog from a layer of silicified volcanic ash in the Darry-Dag near
the town of Dzhulfa.
The marine (Middle Oligocene) Maikop beds of Cis- and Transcaucasia
contain abundant remains of cod and herring. Occurrences of
Halitherium cf. schinzi Kaup. and other, unidentified, sirenian
species (possibly the ancestors of dugongs) are known from the Chiaturi
manganese basin of Georgia, on the southern coast of the Black Sea, and
from the Maikop beds in the Akhzy-Khazry area of the Apsheron Peninsula
(Bogachev, 1938c). The whales Zeuglodon (Microzeuglodon)
caucasicum Lyd. and Iniops caucasicum Lyd. have beendescribed
from beds that are probably the same age (Lydekker, 1892).
Remains of Microzeuglodon aff. caucasicum Lyd. have also
been described from the beds eroded by the Sumgait rivulet 20 km north of
Baku (Ryabinin, 1938).
Occurrences of the whale genera Microzeuglodon, Iniops,
Delphinus (sensu lato) and Zeuglodon have been recorded (Bogachev,
1938c, 1939a) from the Maikop beds of the Apsheron Peninsula.
The same Maikop beds along the Sumgait rivulet contain fossil leaves of
the evergreens Combretaceae, Sapotaceae, Ternstemiaceae, with an
admixture of tropical conifers.
(27)
| | И?
FIGURES 7. Bones of hoofed mammals from the Oligocene of Вепага
1-4 —Pmg,and os lunatum of Benaratherium callistrati Gab.; 2—M*of Aceratherium
cf. filholi Osb.; 3 —metatarsus of Schizotherium chuéua; 5 — fragment of scapula of small
artiodactyl
The abundance of marine mammals and the dense land vegetation attest
to the earlier existence of tropical conditions on the island of Caucasus and
of Sargasso-type lagoons on its southern coast. Observations on the
27 southern slopes of the Taurus Mountains indicate that Asia Minor was also
covered with such forms as podocarpus, myrica, oak, cinnamon,
andromeda and eucalyptus in the Oligocene.
A burial containing Oligocene land mammals was discovered by the
geologist М.Е. Khuchua in 1948 at Benara in the Akhaltsikhe region of
southern Georgia. A few poorly preserved bone fragments and teeth were
found in upper multi-colored sandstone, conformably overlying the lignite
beds (Figure 7). The mode of preservation in situ was inadequately studied.
1704 23
Small bone fragments occur mainly in the concretions made of hard
sandstone. The material was identified and described by Gabuniya (1951а, b,
1953, 1955a, b) as rodent and ungulate:
Rodentia
Fam, gen.
Perissodactyla
Schizotherium chuéua Gab,
Benaratherium callistrati Gab.
Artiodactyla
Anthracotherium sp.
Lophiomeryx benarensis Gab,
28 Gabuniya has correlated the Benara locality with the Oligocene
phosphorite beds of Quercy in France and with the Middle Oligocene
localities of Mongolia and Kazakhstan, where bones of Schizotherium
and Lophiomeryx are known from the Upper Oligocene.
The fauna from the vicinity of Akhaltsykh confirms the paleobotanical
evidence of the land connections which existed, with interruptions, between
the Caucasus and central Asia and between the Caucasus and Western
Europe in the Oligocene.
MIOCENE
In the Lower Miocene the island of Caucasus extended from the vicinity
of Anapa in the west to the upper reaches of the Samur River in the east.
The area of the Dzirul crystalline massif was also an island of smaller
dimensions. The region of Trialeti-Akhaltsikhe and Borzhomi was covered
by sea. Sands were deposited in the sea, although toward the close of the
Lower Miocene the area became dry land (Zhizhchenko, Kolesnikov and
Eberzin, 1940). Only marine mammals are known from the Lower Miocene
beds.
Bones of the seal Phoca sp., resembling P. vindobonensis Toula
(Bogachev, 1927b), occur in the Miocene beds at Lok-Batan, south of Baku.
Remains of Phoca sp. have also been reported from the limestones of
the Kilyazinskaya spit north of Baku, and of the whale Cetotherium
meyeri Brandt from Lok-Batan andthe village of Dzhorat onthe Apsheron
Peninsula (Bogachev, 1938c).
29 The geography of the island of Caucasus at the beginning of the Middle
Miocene, in the so-called Helvetian stage, was essentially the same as in
the Lower Miocene. The sea, in which Tarkhan and Chokrak sediments
accumulated, coveredthe entire area from the northern coast of the island
_of Caucasus to Rostov in the northwest, and Stepnoi (formerly Elista) in
the northeast. At that time the landmass of Caucasia Minor probably
appeared south of Surami island, and the Caucasian island increased in size.
The vegetation of the islands in the Middle Miocene, as shown by the incidence
of fossils in Cis- and Transcaucasia, retained its subtropical character
(Palibin, 1936) though witha small admixture of deciduous trees (Castanea
sativa)andconifers (Pinus neptuni)ofnorthernorigin. In the Middle
24
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25
30
Miocene, in Chokrak time, the Caucasian land gradually became higher and,
in the process of growth, became a peninsula of the southwest Asia massif
(Figure 8).
The sediments of Chokrak age in Ciscaucasia contain abundant remains
of terrestrial mammals. ‘
Central Ciscaucasia
Belomechetskaya. This locality, discovered by A.V. Рапоу in
1926, is near the Cossack village of Belomechetskaya, on the right bank
of the Kuban River, north of Cherkessk (Figure 1).
In this region of the plateau, the Chokrak beds, 60 m thick, are mostly
coarse-grained sandstones plus a combination of sand, clay and calcareous
sediments. In a number of places, sandy bone-bearing lenses have been
exposed in the Miocene hills by the eroding action of the Recent Kuban
River approximately 50 m above the present water level. These were
probably formed in a river which cut through the Chokrak beds eroding an
ancient accumulation of skeletons in a lake or a marsh. The bone-bearing
lense at the southern outskirts of Belomechetskaya is made up of gravelly
quartz sand, with grains measuring 0.5-1.0 mm in diameter, slightly
cemented by silt. Parts of the outcrop are iron-stained in vertical and
diagonal bands. There is no pronounced bedding, which, however, may be
a result of metamorphism.
Areas of silty sandstone, more strongly cemented and of grayish color,
occur within the mass of the gravelly quartz sand. The silty sand indicates
changes in the river current regime. The exposed thickness of the bone-
bearing layer at this locality is 3.5-4.0 m. It is overlain by a limestone
layer 30-40 cm thick. This layer probably corresponds to a brief marine
transgression. Higher in the section the sands alternate with limestones.
In places the sands are covered by Quaternary loams (Figure 9).
The remains of land and marine animals (bones and bone fragments,
fragments of skulls, isolated teeth) are scattered throughout the layer.
They occur in the gravelly sand and more often in the bluish, cemented,
silty and iron-stained sand. The surface of the bones is pale yellow, light
brown or dark brown in color. In fresh fracture the diaphyses are light
brown, the cracks and the pores of the epiphyses are stained with ferric
or, more rarely, ferrous oxide. The tooth dentine is also stained; the
enamel is glossy and dark blue, or sometimes black at the base of the
crown. The old factures on the long bones were rounded by water and the
cavities in the bones were filled with sand.
No tooth marks of predators were found on the bones. It is difficult
to determine why and in what season of the year the animals died. It seems
doubtful that death was caused by catastrophe. The variety and the
fragmentation of the bones, and the abundance of teeth of small herbivores
which occur sometimes in horizontal, sometimes in vertical positions
within the sediment, indicate several cycles of redeposition. The death
assemblage contains both young and old herbivores and carnivores.
Remains of the swamp mastodon (Platybelodon) consist of a fragment
of the facial part of the skull and mandibles which belonged to one adult
and one young individual.
26
31
FIGURE 9. Bone-bearing bed at Belomechetskaya
Photograph by author, 1950
The bone material at this locality includes both terrestrial and marine
mammals, and the aquatic turtle Trionyx. The mode of preservation of
all the bones is very much the same. This indicates the redeposition of the
bones in alluvium. The bones were probably washed from the skeleton-
bearing lenses of Oligocene-Miocene age by rivers or, less likely, by sea
waves. In this process the bones became sorted and redistributed in the
sediments of the river bed. The first burial of the bodies of land animals,
as indicated by the iron stain on the bones, took place in a basin of standing
fresh water, probably on the floodplain. Thus, it is evident that the
existence of the Belomechetskaya ''fauna'’ antedates the deposition of most
of the animals in the fossiliferous layer, although some few of the "fauna'!
undoubtedly died during the time of deposition. In this process the bodies
of land animals carried by the river and the floating bodies of marine
mammals entering the river mouth were macerated and their remains
incorporated into the sediment.
27
32
The collections of PIN, ZIN and the Institute of Paleontology of the
Georgian Academy of Sciences include the species and groups of mammals
given in Table 1.
TABLE 1, Species and number of mammal bones from the Middle Miocene beds at Belomechetskaya*
Number Number
of bones of bones
Camivora Anchitherium aurelianense Cuv, +
Amphicyon Se Dicerorhinus caucasicus Boris. 172
oe Ом Chalicotheriidae gen. 3
Нуаепа Sp, vee ee eee r cece eevee 1
Раза WAUS SD). - Weemowel te Meeeel une n ei alleial +12
Tubulidentata но
Orycteropust о ОИ Kubanochoerus robustus Gab, 1
Hippopotamidae (?) gen. mie сы 1
р Micromeryx flourensianus Lartet 7
Rodentia :
Dicrocerus elegans Lartet 3
Palaeocricetus caucasicus Arg, Paratragocerus caucasicus Soc, 8
Eotragus cf, martinianus Lartet 12
Proboscidea Hypsodontus miocenicus Soc, 1
Platybelodon danovi Boris...... я
MiaIS OIG ON бр еее
Perissodactyla 34
Paranchitherium karpinskii Boris
* Material identified with the aid of Borisyak (1928a, 1943), Argiropulo (1938, 1940c), Sokolov (1949),
Gabuniya (19554, 1956a) and unpublished data of К.К. Flerov and of the author,
The collections also include fish and aquatic turtle (Trionyx). The
total number of known Miocene mammals from the Belomechetskaya is
approximately 20, although the taxonomic identity of some specimens
remains unknown (Figures 10-15).
Paranehitherium, Anchitherium, Rhinoceros and
Platybelodon were considered stratigraphic index fossils by Borisyak
(1937, 1938b). The degree of specialization of Paranchitherium from
Belomechetskaya is the same as that of Parahippus from the Miocene
of North America. Anchitherium, which is probably an earlier form,
is characteristic of the Miocene of Eurasia.
The Belomechetskaya rhinoceros was considered by Borisyak as one
of the most primitive Dicerorhininae. The dentition shows both primitive
and specialized features. Possibly it should be regarded as the oldest
representative of Rhinoceros schleiermacheri.
A peculiar mastodon, Platybelodon, is closest to the genera
Phiomia and Palaeomastodon of the Oligocene of Africa, which
may possibly be the ancestors of the long-snouted mastodon. The lower
tusks of Platybelodon, flattened dorsoventrally and firmly joined,
form an elongated scoop similar in shape to the tusks of Amebelodon
fricki Barb. from the Pliocene of Nebraska. According to Borisyak (1928b)
1704 28
33
34
и ae я
FIGURE 10, Bones of carnivores from Belomechetskaya
1, 2—Hyaena sp.; 3—Amphicyon фр.
7
and Osborn (1936) these mastodons had no trunk. They lived on the shores
of lakes and river backwaters, digging with their flattened tusks for the
succulent water plants and roots which were their food.
Small Dicrocerus, Micromeryx and Eotragus are the main
index fossils of the Middle and Upper Miocene of Eurasia. In general the
fossiliferous layer at Belomechetskaya contains mammals of stratigraphically
different ages. In view of this, it is difficult to identify the index species
of the Middle Miocene faunal complex.
The peculiar features of the Belomechetskaya "Рампа" do not permit
an easy comparison with other Miocene faunas of Eurasia and therefore
its zoogeographical relationships are not clear. However, since the
autochthonous development of such a fauna on the small landmass of the
Caucasus is unlikely, ап earlier land connection between the Caucasus
and the continental landmass in the south can be regarded as certain.
A reconstruction of the Caucasian landforms of the time when the animals
preserved at Belomechetskaya lived would include mountain ridges and
broad valleys with forests, bushland, lakes and quiet rivers as part of the
landscape. The backwaters of those rivers and lakes were probably the
feeding grounds of Platybelodon.
Fossil insects and plants occur in the Middle Miocene beds which contain
marine mollusks. These beds belong to the Karagan and Konka stages.
FIGURE 11, Skull and tooth of Platybelodon danovi (from Borisyak, 1928b)
Poplars (Populus mutabilis), Sapindus fulcifolius and
evergreen cinnamons (Cinnamomum scheuchzeri) occur ш the
Karagan beds of Dagestan. The sandy-clayey littoral sediments of the
Spanidontella sea, southwest of Derbent, contain leaves of evergreen species
characteristic of the Upper Oligocene and Lower Miocene of Europe:
Myrica hakeaefolia, Laurus primigenia, Ardisia cf.
oceanica, Diospyros paradisiaca and other forms. The fauna of
marine mollusks indicates that the Miocene sea became progressively less
30
35
FIGURE 12, Skull and tooth of Paranchitherium karpinskii (from Borisyak, 1937)
saline due to its isolation from the Tortonian basin of the Balkan region,
that the climate became more humid and that the rivers carried more water
(Zhizhchenko, Kolesnikov and Eberzin, 1940).
The richest ''fauna'' of insects (some 90 identified species) was
discovered in bedded marls of the Karagan stage west of Stavropol. The
insects were probably carried by streams into the shallow lagoons of the
northern coast and buried in the silty sediment. The following aquatic
insects are common: mayflies (Ephemeroptera), dragonflies (Odonata),
true bugs (Gerridae), water beetles (Dytiscidae, Hydrophilidae), caddis
flies (Trichoptera) and mosquitos (Diptera). Among the terrestrial
phytophagous, saprophagous, predaceous and parasitic forms are
Orthoptera, Hemiptera, aphids, cockroaches, termites, cicadas, butterflies
(Lepidoptera), Diptera, Hymenoptera and beetles (Rodendorf, 1939).
This assemblage suggests a subtropical climate and profuse grassy
vegetation in the Caucasus at the end of the Middle Miocene.
The numerous plant fossils from the Krynka River valley in the Donets
Basin (Krishtofovich, 1930) indicate that the vegetation on the northern
shores of the Konka basin was of the temperate type and strikingly different
from the subtropical flora of the Caucasus in the Neogene.
In the Upper Miocene the seas still covered the southern Ukraine, the
Crimea, Ciscaucasia, most of the Transcaspian land and possibly Iran.
31
36
37
FIGURE 13. Upper incisors of giant FIGURE 14, Canine of an
perissodactyl from Belomechetskaya unknown artiodactyl
(Hippopotamidae? ) from
Belomechetskaya
The fauna of the Sarmatian sea reflects a further decrease in salinity
and shows considerable variability in facies and in stratigraphic age.
In the Lower Sarmatian the Caucasian landmass grew southward. The
thickness of the Sarmatian sediments indicates that the relief of Japhethida
and of the coastal areas was essentially the same as in the Middle Miocene.
The areas of maximum uplift in the Lower Sarmatian were in the Guri ridge
and in Khevsuretia, and in the Middle Sarmatian, in inner Dagestan. The
northwestern coast remained a plain. The sea in this area became shallower
due to the uplift of the Stavropol massif.
In the Upper Sarmatian the sea receded from the northern part of the
Caspian and from most of the Transcaspian lands, leaving the Stavropol
Plateau as a peninsula projecting into the Ciscaucasian strait. To the south,
a wide isthmus extended from Kutaisi to Telavi connecting Japhethida with
the Caucasia Minor landmass, and in eastern Transcaucasia a narrow Kura
bay formed (Zhizhchenko, Kolesnikov and Eberzin, 1940).
The landscapes of the Caucasus in Sarmatian time can be restored with
a high degree of certainty from the available plant and animal fossil material.
The littoral marine flora consisted of species of Fucus and
Cystoseira, which occur as fossils in the valleys of the Sunzha and the
Sulak on the Malokabardinskii ridge.
Seals (Phocidae) andsmalltoothless whales ofthe genus Cetotherium
were abundant in the Sarmatian sea. Bones of Cetotherium have been
found in the Sarmatian coquinas and silts near Derbent, in Makhachkala,
32
and in the blue Sarmatian clays near Maikop (Spasskii, 1939). Bones of seal
have been found in the white marine sands near Stavropol, in the blue clays
near Maikop and Goryachi Klyuch and in a number of other places. The
Stavropol seal bones were found with bones of terrestrial Tertiary mammals
— undoubtedly species which lived in a warm climate.
FIGURE 15,
1 — jaw of Kubanochoerus robustus (from Gabuniya, 1955a); horn axes of: 2 —Paratragocerus
caucasicus; 3—Eotragus cf, martinianus; 4—teeth of Hypsodontus miocenicus (from
Sokolov, 1949)
The terrestrial flora of the Caucasus in the Sarmatian is represented
by coniferous, evergreen and deciduous Species. Remains of spruce
33
38
(Abies sp.)*, pines (Pinus sp.) and sequoia (Sequoia sp.) occur in
the exposures along the Supsa River. Magnolia, laurel (Laurus
primigenius), cinnamon (Cinnamomum polymorphum), elm,
willow and pear trees have been identified from the Kakhetian Range and the
Trans-Kuban Plain.
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FIGURE 16, Land formations in the Upper Miocene (Upper Sarmatian)
(From Kolesnikov; see Zhizhchenko and others, 1940, )
Dots indicate main occurrences of land vertebrates
At some of the Middle Sarmatian localities in eastern Georgia as many
as 30 arboreal species have been collected. Of these, up to 70% are
deciduous and include the Tertiary species of hornbeam (Carpinus), oak
(Quercus), willow (Salix), maple (Acer), chestnut (Castanea),
zelkova (Zelkova ungeri), and shrubs of Indian hemp (Apocynum),
bladder senna (Colutea salteri) and other deciduous species in addition
to the evergreen laurel, cinnamon, myrica and magnolia. According to
Palibin (1936) the evergreen species of the flora ''which covered the entire
surface of Japhethida'' were the last Sarmatian representatives of the
subtropical and tropical flora which flourished on the Caucasus from the
beginning of the Tertiary. This first step in the borealization of the flora
contributes to an understanding of the later evolution of the fauna. Ina
comparison of the Caucasian flora with the Upper Miocene flora of Asia
Minor, Grossgeim (1936) found that they are very similar, both being
characterized by a mixture of boreal and subtropical elements.
The taphonomy of these dead plant assemblages is significant for the
ecologist or faunist in the study of altitudinal zonation of flora which
“ [There is a discrepancy in the Russian text between the common and the Latin name here, The latter
corresponds to the true firs, whereas the Russian gives the name "spruce, "]
34
39
might have developed in the presence of the mountainous relief that existed
in the Middle Miocene. Such a zonation might explain some of the instances of
typically ''mixed'' Miocene flora.
Hipparion fauna appeared in the Caucasus in Sarmatian time, having
migrated from the south along the wide Transcaucasian Isthmus (Figure 16)
it was probably first discovered in Transcaucasia by Ryabinin (1913).
Ryabinin found bones and teeth of Hipparion gracile Kaup and
tooth fragments of Rhinocerotidae in the Kyasaman site and on the left bank
of the Iora River. At this locality the vertebrates occur in red clays with
sandstone intercalations (which also contain algae (Chara cf. escheri)
and mollusks (Planorbis sp.))andthe same mammalian genera were
recorded by Ryabinin from gypsiferous clays underlying the Akchagyl beds
in outcrops in the Katsakhuris-Kedy ridge.
In 1913, the geologist Dombrovskii, working further south on the right
bank of the lora River, found in the cliffs of the ЕПуаг-Опва and Palan-
Tikyan ridges a rich locality of Tertiary mammals, known as the El'dar
locality (1914).
i aliete sy llama ey IP bene @ el wis
El'dar. According to the descriptions of Dombrovskii (1914), and
Andrianov and Larin (1935), the bone-bearing bed can betraced over 6 km
from the gorge which connects the El'dar Steppe with the Iora River
floodplain. From west to east the bed grades from a shell conglomerate
to sandstone, then to limestone, and again to sandstone which grades into
thick upper Sarmatian clays where the bed eventually peters out. The bones
(parts of skeletons) occur in pockets, at intervals of 10, 20 and a few
hundred meters. Ripple marks occur on the top surface of the bone-bearing
sandstone.
The lithology of the bed attests to the existence of a low coast and muddy
bay. The bodies of animals were transported into the bay by weak currents
and shore waves and were buried in the sandy-clayey bottom sediment. The
investigators believe that mass mortality in the animals was caused by
steppe fires, earthquakes and mudflows.
According to our observations, all the complete bones, including lower
jaws and ribs, were flattened by the pressure of the overlying beds. Some
bones were broken in the process of burial; as a consequence, the cavities
became filled with sand and silt which preserved the original shape of the
bone.
The surface of the bones is chestnut or chocolate in color; in fresh
fracture, the colors are grayish. Permineralization and diagenesis of the
bone material were not significant. Preserved coprolites were also found.
Bones abraded bythe surf are rare, and in many cases limbs were
preserved with bones intact. In general, a number of features support the
conclusions of earlier authors regarding the nature of the burial and the
mode of preservation.
Table 2 (Figures 17, 18) lists the species from the Moscow and Baku
collections identified by Bogachev (1927a), Alekseev (1930), Borisyak and
Belyaeva (1948).
35
41
TABLE 2, Species and number of mammal bones from the upper Miocene beds at El'dar
Number
of bones
Primates Artiodactyla
Bas, PEW swendenoaysdaree р ИР 1 Sus erymanthius Roth, et Wagn, 5
Achtiaria borissiakii Alex. ....
Carnivora Camelopardalis (Helladotheri- 69
с sans : О akericka tae БАРОН ЯОЧОЙЬ a
be ha eee ds Regs RT ISS}: АИ Tragocerus aff, leskevitschi
Hyaena cf, eximia Gaudry....... 6 Е
BOEES. ее. А. 66 =
| ар осетр. ИМО ЗЕ ое 159
Proboscidea
TERA OC ети зато eens сы
Mastodon longirostris Kaup..... GE Zie а Бр. gh: с оо “baal omy oi 6
МЕ piemimemiiCi Gadi ysis weet ean element 24
Mastodon sp. a «+> = aise wee Pinnipedia
j р С : f 3
Dinotherium giganteum Kaup,.. Phoca sp eee ТО 5
Perissodactyla Gb
: . : 164
тратой Е ОР Ри Е у оО Ооо ооо ов еовос
спеши ииияя 955 обо боньс 3 2 : 208
г г сезоне Spe) И бе а Же =
Aceratherium transcaucasicum
Во шо Goh laa Saco. 22049 Оно he 6
Dicerorhinus aff, orientalis
БО И. te seh ею Ban 0 65
The El'dar collection may be regarded as a part of the Upper Miocene
faunal complex of Transcaucasia, since there are no signs of redeposition
of the bones and most of the species lived at the same time. The most
abundant species, as far аз the collections show, were Hipparion, ibex,
rhinoceros and giraffe.
Udabno. This sizable locality containing Hipparion fauna was
discovered in 1931 by N.A. Gedroits near the village of Udabno, on the
Kura-lora water divide of the Garedzhiiskaya Steppe. The locality was later
excavated by the Museum of Georgia.
The fossils occur in Sarmatian clays overlain by sandstones of the Shiraki
formation. This locality and the El'dar locality are similar to each other,
as are the lists of species from each. The material is in the collections
of PIN and the Museum of Georgia, and was identified by Burchak-
Abramovich and Gabashvili (1945, 1950), Tsereteli (1942), and Borisyak
and Belyaeva (1948):
Primates Proboscidea
Udabnopithecus garedziensis Burtsch, Mastodon sp.
et Gab, Dinotherium sp.
Carnivora
Hyaena sp, Perissodactyla
Rodentia Hipparion gracile Kaup
ine Aceratherium $
Hystricidae gen, Be
36
(40)
FIGURE 17, Remains of carnivores and hoofed mammals from El'dar
1, 3—skull of Hyaena sp.; 2—coprolite of hyena; 4, 5 — tarsal joint and horn axis of Tragocerus
sp.; 6, 7—horn axis and jaw of Gazella sp.
Artiodactyla
- у Achtiaria sp.
Sus sp. Tragocerus sp.
Cervidae gen. Gazella sp,
The occurrence of teeth of alarge homonid ape (Udabnopithecus
garedziensis) is of particular interest in connection with studies of
the origin of man (Figure 19).
FIGURE 18, Remains of carnivores and hoofed mammals from El'dar
1— jaw of Crocuta eldarica; 2—incisor of Aceratherium transcaucasicum;
3-5 — teeth of Tragocerus leskevitschi (from Bogachey, 1927b)
Arkneti. This locality is 1.5 km east of the village of Arkneti in
South Ossetia. It is situated on what was the western part of the
Transcaucasian bay of the Sarmatian sea. Fragmented bones of mammals
occur in the yellowish gray loams, 1.5-2.0 m thick. Well-preserved bones,
some of them joined, and skulls occur in a bone-bearing lense 3 m long
and 50-70 cm thick. The artiodactyls and perissodactyls have been
identified by Gabuniya (1952b, 1955c):
Perissodactyla Tragocerus ex gr, leskevitschi Boris,
Tragocerus sp,
Gazella cf. gaudryi Schlos.
Gazella sp.
Giraffidae gen.
Sus sp. Phronetragus arknethensis Gab.
Eostylocerus sp,
Dicrocerus salomeae Gab,
Hipparion sp. (cf. garedzicum п. sp.)
Artiodactyla
38
12
FIGURE 19. Teethof Udabnopithecus garedziensis
1-4 —Pm*. anterior, outer, inner surfaces (x12), upper
surface (x 24); 5-7 —M*(x14) anterior, outer and upper
surfaces (from Burchak- Abramovich and Gabashvili, 1950)
The age of the ''fauna,'' according to Gabuniya, is very close to the age
of the Udabno ''fauna''; it may be dated as Upper Sarmatian (Meotian).
The existence of large valleys covered with savannah and tropical forest
vegetation on the piedmont can be inferred from the composition of the
Sarmatian fauna. Streams originating in the Caucasus Range were evidently
sufficiently powerful to transport the bodies of mastodons and rhinoceroses
into the coastal bays of the Sarmatian sea.
As а whole, the Sarmatian faunal complex supports a picture of the
landscape drawn from the known paleontological data. The plant fossils
indicate not only the ''mixed'' character of the Sarmatian flora, but also
the high degree of its differentiation.
Stravropol area
The Hipparion faunal complex occurs in the Sarmatian of the
Ciscaucasus. The fauna migrated along the coasts and over the plateaus
and ridges of the central part of the peninsula. Hipparion sp. and
Dinotherium giganteum Kaup are known from the Middle Sarmatian
limestones near the village of Burlatskoe (Khomenko, 1913a). A small
giraffe, Camelopardalis parva Weith., is known from the Upper
Sarmatian beds near the village of Blagodarnoe in the Stavropol area
(Pavlova, 1933a).
Mount Kutsai. A fairly rich locality was discovered by A. A. Ivanchin-
Pisarev in 1915 near the village of Petrovskoe, northeast of Stavropol.
Like other Transcaucasian burials of Sarmatian age, it contains both
marine and terrestrial vertebrates.
39
43
The bone-bearing beds of Mount Kutsai are made of sands and clays
(Ivanov, 1916). These are underlain by sandy, plastic clays of Middle
Sarmatian age; soft limey sandstones overlie the fossiliferous sequence.
The fossils occur in medium-grained marine quartz sands. The material
is poorly preserved and consists mostly of individual teeth, fragments of
vertebrae, metapodia and phalanges. Bones of seal and whale occur in the
lower 2-3 m of the Upper Sarmatian clays and sands. The bones of
terrestrial forms occur somewhat higher, although still within the lower
3-5 m of the clayey-sandy beds.
Ivanov's (1916) identification of the terrestrial mammals reveals the
presence of the Hipparion faunal complex:
Perissodactyla Gazella sp,
Hipparion gracile Kaup. se ВЕТ.
Hipparion sp,
Pinnipedia
Aceratherium sp, 4
Rhinoceros sp, Phoca cf, pontica Eichw,
Artiodactyla Cetacea
Listriodon sp, Cetotherium priscum Brandt,
Sus sp. Cetotherium (?) sp.
Remains of birds, turtles and fishes also occur at this locality.
Individual occurrences of the bones of rhinoceros (Dicerorhinus
and Aceratherium) are known from the greenish silts and sands near
Maikop and from the marine sands near Beshpagir and Stavropol. Asa
whole, the composition of the fauna of Ciscaucasia in the Sarmatian was
very Similar to that of the Transcaucasian fauna.
In order to identify the origin of the Hipparion faunal complex which
appeared in the Caucasus, we should compare the Sarmatian ''fauna"' of the
Caucasus with other Miocene ''faunas'' of Eurasia. The following groups are of
interest: the Miocene ''faunas'' of Punjab — in the southern foothills of the
Himalayas in the Salt Range in the Indus River valley, and in the Siwalik
Range in the Indus and Ganges basins; the Middle Sarmatian ''fauna'' of the
Crimea; the Upper Miocene ''fauna" of Iran from the eastern shore of Lake
Urmia; and other occurrences in Asia Minor and the Balkans.
The ''faunas" of the Siwalik and the Salt ranges occur in continental
deposits upto 7,000 тп thick (Colbert, 1935). Over 288 species of mammals
are known from these Upper Miocene to Lower Pleistocene formations,
which evolved from the erosion of the newly uplifted mountain range. At
this time detrital material was deposited on a land surface, sometimes
partially covered by freshwater bodies.
The Russian paleontologists and geologists are of the opinion, taken
probably from Osborn (1921), that the Hipparion "fauna" of southern
Asia (Siwalik Range) migrated via the Caucasus into Eastern Europe.
According to Andrusov (1918) the [Hipparion] fauna of the Pikermi
type appeared in Europe from the east, following two routes: one along
the northern coast of the gradually shrinking Sarmatian sea and one through
Asia Minor.
44
Andrusov shows that the development of the Hipparion fauna was not
complete in the Middle Sarmatian but continued through the Upper Sarmatian,
reaching its maximum level in the Meotian. In southeastern Europe the
fauna probably became extinct in the Lower Pontian.
Borisyak (1928a, p.376) assumed that the Hipparion fauna migrated
to Europe from Asia via the Caucasus and the Black Sea landmass.
According to Bogachev (19384, рр. 36-37), ''the African-Siwalik fauna
spread through Iran and Transcaucasia onto the growing Main Transcaucasian
(sic!) Range, and from there, the migration continued across the Stavropol
Plateau onto the Ukrainian Steppes. The stages of migration are marked
by fossils which occur along the migration route. "
However, Colbert's lists of the Upper Miocene fauna of Kamlial and
Lower Siwalik show that there is practically no resemblance to the
Sarmatian fauna of the Caucasus.
Nearly 85 species of primates, rodents, carnivores, proboscideans,
perissodactyls (Hipparion, rhinoceros) and artiodactyls (swine, tragulids,
deer, giraffe), were noted to 1935 in the upper beds of the Lower Siwalik
in the Salt area (Chinji). It is Significant that no Cavicornia occur in these
beds. The equids, rhinoceroses without horns, giraffids and primates are
found in both the Caucasian and the Lower Siwalik faunas.
The Sevastopol ''fauna'' in the Crimea, assigned to the Middle Miocene
age, consists of the following forms (Borisyak and Belyaeva, 1948):
Carnivora Artiodactyla
Ictitherium tauricum Boris. Achtiaria expectans Boris,
Tragocerus leskevitschi Boris,
Perissodactyla Tragocerus sp.
Gazella sp.
Hipparion gracile var, sebastopolianum
Boris
Aceratherium zernovi Boris,
A, zernovi var, asiaticum Boris, Testudo sp.
Trionyx sp.
Reptilia
The Stavropol faunal complex was undoubtedly very similar to the El'dar
complex, although the zoogeographical relationships between the two are
not clear.
Recent geological data indicate that the Hipparion complex could only
have reached the Crimea from the southwest at the end of the Middle Miocene
or in the Lower Sarmatian. Later migrations can be ruled out, since the
Caucasus landmass was probably cut off from the Russian platform and from
the Crimea by the sea which existed until Pliocene time (Andrusov, 1918, 1926;
Muratov, 1951).
The ''fauna'' occurring in the clays and sands at the southern slope of
Mount Sahand near Maragheh is of particular interest in the reconstruction
of the Miocene landscapes and the faunal ties between Asia and the Caucasus.
According to the identification of Rodler and Weithofer (1890),
Mecquenem (1924), Bogachev (1928), and Burchak-Abramovich (1952b),
the following species are represented in the Maragheh ''fauna’':
41
Primates
Mesopithecus orientalis Kittl
M, pentelici Gaudry
Carnivora
Hyaenarctos maraghanus Меса,
Ictitherium hipparionum Gaudry
I, robustum Gaudry
Hyaena eximia Wagner
Meles polaki Kittl
M. maraghanus Kittl
Felis brevirostris Croiz, et Job.
F, attica Wagner
Machairodus aphanistus Kaup
М. orientalis Kittl
45
Tubulidentata
Oryteropus gaudryi Major
Proboscidea
Mastodon pentelici Gaudry
Perissodactyla
Aceratherium persia Pohlig
Rhinoceros morgani Mecq.
Chalicotherium pentelici Gaudry
Hipparion mediterraneum Hensel
Artiodactyla
Sus (Microstonyx)erymanthius Roth.
et Wagner
Helladotherium gaudry Mecq.
Alcicephalus neumayri Rodl, et Weith.
Camelopardalis attica Gaudry
Urmiabos azerbaidzanicus Burtsch.
Gazella gaudryi Schlosser
С. brevicornis Gaudry
С. capricornis Rodl, et Weith.
Palaeonyx pallasi Gaudry
Protoryx carolinae Major
Antilopinae gen. ?.
Tragocerus rugosifron Schlos.
Protragelaphus scozesi Dames.
Tragelaphus hontom-schindleri Rodl.
et Weith.
Helicophora rotundocornis Weith.
Oiceros rothi Wagner
О. atropatanes Rodl, et Weith.
O. boulei Mecq.
Bones of ostrich (Struthio sp.) and Urmiornis maraghanus
Meq. have also been recorded at the locality.
The large number of species in the Maragheh ''fauna'’ attests to a high
degree of universality of the death assemblage.
Although the taphonomy
is not known in detail, it seems likely that the death of the Upper Miocene
animals can be attributed to a common cause. This might have been toxic
gases (fumaroles), volcanic ash deposits or mudflows. According to Pohlig
(1886), the Maragheh ''fauna'' was buried near the shores of Lake Urmia,
the water level of which was much higher in the Miocene. The sedimentary
facies at the locality are similar to the Pikermi locality near Athens and
the Val d'Arno locality near Florence.
The great variability of the
assemblage is reflected in the occurrence of representatives of different
ecological habitats: arboreal (e.g., Mesopithecus) and savannah and
steppe (e.g., giraffes, gazelles, ostriches), and of different feeding types:
carnivores, omnivores, herbivores. A predominance of savannah forms
over subtropical forest forms indicates that the landscape was of a mixed
savannah-tropical forest type, like that in the northern parts of the Iranian
Plateau in the Miocene.
The Mardagheh ''fauna'' has some ''African'' elements in it. It is fairly
close to, though not identical with, the older Sarmatian fauna of
Transcaucasia.
According to Stahl (1907) and Oswald (1915-1916), the interior of Iran
and Anatolia became dry land in the Upper Miocene. Salt and gypsum
precipitated in the relicts of the Miocene seas.
Bogachev and Shishkina
(1915) discussed the importance of these geological processes in the
formation of bone-bearing deposits. According to these authors, the climate
of the country became arid in the Middle Miocene following the uplift and
folding which subdivided the marine basin into a number of saline lagoons.
42
This notion of an arid desert climate is not confirmed by ecological
analysis of the Maragheh mammals. It is more likely that tropical forests
existed on the shores of the relict basins, although the climate as a whole
was dry and hot.
The Maragheh faunal complex was probably characteristic of all of
southwest Asia. This is confirmed by geological data on the paleolandforms
of Asia Minor (Furon, 1955).
Occurrences of Upper Miocene mammals in Asia Minor are known from
Stambul, upper Gediz, Mugla, Galatia and Cappadocia. The collections
comprise carnivores (Ichtitherium, Martes, Machairodus),
proboscideans (mastodon), perissodactyls (Hipparion and rhinoceros)
and artiodactyls (giraffe, gazelle, antelope). A similar fauna’ is found on
Samos Island and in the Balkans, near Athens (Pikermi). The volcanic
eruptions in Cappadocia, accompanied by ashfalls and mudflows, caused
death among animals and subsequent burial in layers of tuff.
Аз a whole, the Upper Miocene faunas of southwest Asia, the Caucasus,
the Crimea and the Balkans are similar to one another. However, the order
of appearance of the Hipparion faunal complex in each area cannot be
established until all the material has been studied, a task beyond the
abilities of one investigator.
At the end of the Miocene, in Meotian and Pontian times, the seas
surrounding the Caucasus became shallower and less saline. The Stavropol
Plateau continued to grow towards the north, the area of the present
Dagestan Mountains extended to the northeast, the eastern Caucasian gulf
almost disappeared; and a narrow strait probably existed intermittently
in the Manych area.
The major Tertiary uplifts of the Caucasian Range had probably ended
by that time, andthe highland faunal complexes proceeded to form under
conditions of high-zonal climates and probable local glaciations.
There is no evidence of any extensive glaciation in the Caucasus in the
Upper Miocene, as hypothesized by Kovalevskii (1936) in his study of the
continental formations at Adzhinour, Nor does the paleontological evidence
support the hypothesis of Kovalevskii and Grossgeim (1936) of extinction
of the tropical flora and fauna in the Caucasus caused by an Upper Miocene
glaciation. Neither is there any evidence that the Gunz and Mindel glaciers
were so large that they could ''plough"'’ and fill the Alazan-Agrichai valley
in eastern Transcaucasia, which, in any case, is probably not that old.
Grossgeim (1936, 1948) believed that in Meotian time the southern
xerophilous flora invaded the Caucasus. However, the fossil record shows
that the only flora known is the mesophilous forest flora in Gurie [Western
Georgia] in which deciduous species (Fagus orientalis, Acer
trilobatum) predominated, but which also included evergreen subtropical
forms (Rhododendron pontic um) and some admixture of conifers
(Taxus grandis). Mesophilous plants were also recorded at Nakhichevan
in the Lower Pliocene rock salt deposit (with paper shales): alder, mazzard,
hop, hornbeam, willow, sedge and reed (Palikin 1936, Bogachev and
Shishkina 1915).
Fossil plants and animals of the highlands are not known from that
period. However, it is possible to assume that, in addition to some endemic
evolution, cold-climate flora and fauna migrated to the Caucasus over the
ranges of the Alpine region.
43
47
The extant forms which evolved in the Upper Miocene are the Asia Minor
hamsters (Mesocricetus), Prometheomys schaposchnikovi
and the wild Caucasian goat.
The Uppermost Miocene terrestrial fauna which inhabited the piedmont
plains is as yet little known and poorly dated. The Hipparion faunal
complex continued its existence on the plains around the Caucasus mountain
system, particularly on those to the southeast.
Rare occurrences of mammals are known in eastern Transcaucasia and
in western Ciscaucasia. One was discovered by N.A. Kudryavtsev in 1935
on a northern spur of the Kakhetian ridge in Transcaucasia.
The stratigraphic position of that locality is not clear, although it has
been tentatively datedas Upper Miocene or Lower Pliocene. According to
Orlov (1936b) the bone-bearing bed is exposed on the left slope of a deep
gorge which cuts through the village of Dzhaparidze (near Tsiteli-Tskharo)
toward the Alazan River. Scattered and broken limbbones, jaws and teeth
of mammals occur in marine (?) clays overlain by continental clays
alternating with layers of fossil soils. The following species are known
from the locality:
Proboscidea Artiodactyla
Fam, реп. Sus sp.
Cervidae gen,
Perissodactyla Giraffidae gen,
Gazella sp,
Hipparion gracile Kaup
Rhinocerotidae gen,
In Ciscaucasia the Upper Miocene mammals were collected by
Ya.M. Eglon in 1940 from the Meotian (?) clays on the right bank of the
Kuban River near Armavir. Individual fragments of permineralized bones
were identified as Hipparion sp., antelope about the size of saiga
(Antilopinae gen.), small beaver (Castoridae gen.), small seal (Phoca
sp.) and saber-tooth (Machairodus sp.).
Fragments of molars of a rodent (Muridae), about the size of a house
mouse, were found by V. Sizov in the Miocene beds of the River Aksai
ravine in the Grozny Region at the boundary of the Upper Sarmatian and
Meotian yellow-green clays. Remains of grasses, freshwater ostracods
and otoliths of gobies (Gobius) were also collected.
The vertebrates from the continental deposits near Novocherkassk, in
the south of the Russian Plain, were tentatively dated Meotian age.
The bodies of animals were probably carried by the paleo-Don into the
Tanais Gulf of the Meotian sea where they were buried in the crossbedded
white sands. The stratigraphic position of the sands is ''between the Pontian
and the eroded Sarmatian'' (Zhizhchenko, Kolesnikov, and Eberzin, 1940).
The sands are up to 12 m thick. Very thin intercalations of bluish grayclay
with diatoms, silicified wood and teeth of proboscideans occur in the sands.
Near the village of Yanov, mollusks (Congeria and Neritina)
and perch bones (Perca sp.) were collected from the sands. The mammals
collected by Lisitsyn and Bykodorov near the villages of Yanov and Popovka
and inthe quarries along Tuzlovka River were identified by Квотепко,
Bogachev and Sokolov (1954) as belonging to four species:
48
Proboscidea Dinotherium gigantheum Kaup
Mastodon borsoni Hays, Artiodactyla
М. ef. tapiroides Cuv, Palaeoryx longicephalus Soc,
Thus the end of the Miocene in the Caucasus was characterized by the
growth of landmasses, the appearance of northern species in the flora and
the development of altitudinal zones of vegetation resulting from cooling of
the climate and mountain-building movements. These processes, takentogether,
promoted the rapid evolution of horses, Cavicornia, ruminants and deer,
and a concomitant decline of giraffids.
PLIOCENE
In the Caspian region, four stratigraphic subdivisions of the Pliocene
can be recognized through the fauna: Pontian, Balakhany, Akchagyl and
Apsheron. In the Black Sea region, there are five subdivisions in the same
period: the Pontian and four subdivisions above it corresponding to the three
upper subdivisions of the Caspian: Cimmerian, Kuyal'nitsk, Gurie and
Chauda.
The outline of the Caucasian Peninsula did not change essentially from
the Upper Miocene to the Lower Pliocene (Lower Pontian). The peninsula
extended into a slightly saline lake-sea, not connected with the ocean
(Andrusov, 1918). Two large open bays cut into the northern coast of the
lake: the Kuban bay inthe northwest and the Terek bay in the northeast.
In the western part of present-day Colchis, the Rion bay extended to the
longitude of Kutaisi and to the east lay the Samur and Kura bays. The thick
Pontian sediments andthe littoral conglomerates (Kolesnikov, Zhizhchenko,
and Eberzin, 1940, p.402, map) indicate that all of these but the Kuban were
very deep. Their depth suggests a considerable uplift in the coastal areas
of the region which today comprises Abkhazia and northern Azerbaidzhan.
The broad land connections with southwest Asia allowed the migration
of southern plants and animals to the Caucasus.
A number of geologists (for example, Sokolov, 1904) have noted that
in Lower Pontian time the climate became cooler and the rivers of the
Russian Plain periodically froze.
Nevertheless, the climate of the Caucasus remained fairly warm in
Lower Pliocene time. The abundant plant remains in the Yergeni beds (50 km
north of Stalingrad, in the Ilovlya River Basin) represent deciduous, warm-
climate flora of the Upper Miocene-Lower Pliocene. The flora consisted
бога Розе АТ щаб тасава. | ube we Cisse, 1 C ars iteamera:
sp., Parrotia persica, and Araliacea (identified by Baranov, 1952).
The occurrence of the Russian pea shrub (Parrotia persica) inthe
Yergeni beds is a good indicatcr of a warm climate in the Lower Pliocene,
since this species only survived through the Pleistocene south of the
Caspian sea, i.e., 10° in latitude farther south.
With the Middle Pontian, the climate in the Caucasus became warmer,
as evidenced by plants collected in stratigraphic sections in the Pontian
sediments at Cape Pitsunda (Mchedlishvili, 1954b).
45
49
At the end of Pontian time, the seas receded from Ciscaucasia and the
water remained in the Caspian region only in the southern part of the basin
(Figure 20). Extensive semideserts probably formed in eastern Ciscaucasia
and eastern Transcaucasia.
Later, in Cimmerian time, the climate became almost tropical, as
indicated by the occurrence of banded iron-ores in the Taman and Kerch
peninsulas and in the foothills of the northern Caucasus. Tropical conditions
can also be inferred from the occurrence of tropical plants in marine
littoral sediments (Mchedlishvili, (1954a) and in volcanic tuffs at Goderdzi
on the Adzhar-Imeretia ridge in Transcaucasia (Palibin, 1936). The plant
remains include Ficus, Araucaria and palms.
35 40 45 50 55 60
= SL ok р. р
Smolensk Т м vagen?
eg [5 Kuibyshey
Sa amarky р»
о У / ait,
[ Kursk < Ре) 9 Kant
a Kiey )
[\ Kharkoy% ra
} : 2 : oe
3 )
Ф Stalingrad «4 >
? - - р
tina TT tn, ond ЧТ my >
“lily lista Sil ets as:
feropo
aa
oe itil то в. é
BLACK Ig, : К N
Se as rozny | —_
и. ul was LLL [Ц ЧЩ | In ‘Tbilisi $.
С i : en in ;23103985%
WW [0 ome Batumi ¢ = БА ce
fesse CEO gh
ыы np ew gr :
25 30 35 40 45 50
FIGURE 20, Cimmerian and Balakhany basins (from Kolesnikov; see Zhizhchenko and others, 1940)
Dots indicate main localities of land vertebrates
The land connection between the Caucasian Peninsula and the Russian
platform was established for the first time in the Cimmerian; this allowed
migration of plant and animal species in both directions.
At that time the fauna of the deserts of central Asia could have migrated
far west to the eastern plains of the Caucasian Isthmus. During the
Cimmerian, the configurations of the Black and Azov seas were similar
to their Recent configurations.
Lower and Middle Pliocene terrestrial vertebrates of the Caucasus occur
primarily in continental deposits.
46
50
Stavropol Plateau
The largest and most complete body of Pliocene material was found
10 km west of Stavropol in a locality of sandy sediments which were exposed
in the Kosyakin quarry. А bone-bearing lens, 7-9m thick and 90m
wide, was formed in a channel of a Pliocene river which cut through the
Sarmatian limestones at the edge of the Stavropol Plateau (Kaspiev, 1939;
Gnilovskoi and Egorov, 1955).
Seven beds can be distinguished in the old alluvium. The lithology of the
beds varies from fine-grained, grayish white sand to gravels with clayey
intercalations and clayey pebbles up to 1.5 cm in diameter (Figures 21, 22).
— 5
06
10
hn
|
Mh
,
FIGURE 21, Stratigraphic section in the Козуа п quarry
a, b — loams with humus; c-g — bone-bearing river sands, some of them
crossbedded. Numbers on the right indicate thickness in meters
Our observations show that the pebbles in the lower part of the section
are made of Sarmatian local rocks, rounded bones of Sarmatian marine
mammals and, more often, Pliocene land mammals.
Bones of animals, fractured and redeposited many times, were retained
in the gravelly sands which were deposited at [current] velocities of
1.3 m/sec (lower bed) and 0.1 m/sec (upper bed).
The bones were preserved in various stages of abrasion, from complete
specimens to rounded pebbles.
Up to 92% of the bones collected in the quarry are free of organic
matter and nearly free of permineralization. The bones resemble friable
marl or chalk. The surface color of the bones is cream or grayish. All the
cracks in the bones of this type are iron-stained. Manganese and iron
dendrites occur rarely in the bones. Thin wavy grooves, the marks of
47
51
Recent plant roots, are common on the bones. Notches of irregular shape,
formed by water erosion and aquatic invertebrates at the time of deposition,
were observed on some bones.
The bones in this group are, for the most part, intact; fractures are
mainly the result of careless collecting during quarrying. The remaining
8% of the bones are gray, and numerous dendrites give the surfaces a stained
appearance. As arule, these bones are more strongly permineralized and
considerably heavier than the ''chalky'' bones. The tooth enamel is always
well preserved; its color is either natural or light pea green or light pink.
Tooth impressions of carnivores are rare: only nine bones show signs
of gnawing on the epiphyses.
FIGURE 22, Kosyakin quarry
Photograph by author, 1952
The occasional vertebrae of Sarmatian whales and seals, derived from
the Sarmatian limestones, are easily distinguishable from Pliocene bones
by the heavy permineralization and grayish brown color,
Variations in the conditions of deposition and in the exposure times on
river banks and spits during redeposition account for the differences in the
state of preservation of the Pliocene bones.
It seems likely that the physical environment of the valley and stream
of the Stavropol paleoriver was similar to the middle parts of the Don, the
Volga and the Ural in Quaternary time. Bodies of dead animals accumulated
in the oxbow lakes, as well as in the mainstream and backwater.
In the process of erosion of older marine sediments, the river carried
the bone material over short distances and redeposited it.
The direction of the stream flow has not yet been established. Perhaps
when the sea level was low and the Stavropol area slightly elevated, the
48
52
53
river source was somewhere on the Russian Plain (e.g., near Yergeni)
rather than on the Caucasus.
In spite of the different ages of the bones at the locality, most of the
Pliocene species can undoubtedly be regarded as one faunal complex.
The fragmented material makes identification difficult, and most of the
species have not yet been described in detail.
A preliminary list of the species is given in Table 3 (collections of PIN
and ZIN: identification by Belyaeva (1940b, 1944), Borisyak and Belyaeva
(1948), Argiropulo (1939b, 1940c), and Vereshchagin (1954)).
TABLE 3, Composition of the Pliocene fauna and number of bones from the Kosyakin quarry
Number Number
of bones of bones
Insectivora Proboscidea
a EmOlCiGiiral < еее Anancus arvernensis Croiz, et Job. 21
BY INN DENN о о бо дБ ооо бое ПЕ © НИ GI. оороваасе в овоб 12
“IDES WAAL Gy в оосоевововосо
Perissodactyla
Carnivora : р
ND) DAWLOM S95 5 oaaaoonocaga00% 3
* Ursus cf, arvernensis Croiz, EL up)pial milo meseaatenale акр 18
Р1пос\ on cf, thenardi Jourdan Tapirus cf, arvernensis Dev, et
САН SOM, OF ono Hola Ao eo BOM обо або во воносбовоа 3
Felis cf. issiodorensis Croiz, Dicerorhinus orientalis Schlos. 72
ЕВ ats ain с Мы Aceratherium cf, incisivum Каир 8
Rhinocerotidae gen, (cf. Chilo-
Lagomorpha В) 55s oon mows ooo oo 646 1
Е ри р Ев", АЙ Mie oo eR eR rae: te
7” :
Ochotonal ch antiqua ... le Artiodactyla
Rodentia Propotamochoerus provincialis
ео Во мен СЕТИ: о и olevian eet 10
2 SUCMCORMDEP 95. соооооььвово Е UO HEA? Re И |
: - Palit Oleleny У рр ли 9
СОВА 95 ро ие в
О ре Ooo, 0 Seog о Е ит a
Giraffidae gen. (cf. Sivatherium)... 2
Сале ИТ афера, и causeway metab 5
Note, Asterisk indicates author's collection of 1952.
In addition to the land mammals given in Table 3, the material includes
birds (Charadrius cf. morinellus L.), aquatic turtles
(Trionychiidae — 14 carapace fragments), the lower jaw of asmall lizard
(Lacertilia), and bones of small anurans (frogs and toads) (Figures 23-25).
The faunal complex of the ancient Stavropol area was well developed.
The Stavropol mastodons, tapirs, rhinoceroses and warthogs lived among
subtropical mesophilous plants. Groves of tugai vegetation near the rivers
probably alternated with open meadows inhabited by moles, hamsters
and pikas.
In the preceding sections we reached certain taphonomic conclusions
and outlined a landscape scheme. These judgments are, to an extent,
49
confirmed by the following phenomena: the predominance of fossils of
animals associated with river valley thickets (rhinoceros, swine; Figures
26 and 27), the occurrence of beaver, tapir and aquatic turtle fossils in
swamps, oxbow lakes and channels, and the iron stain which appears on
these fossils. The presence of small desmans in the Pliocene river indicates
FIGURE 23, Bones of smaller vertebrates from the Kosyakin quarry
1 — jaw of lizard (Lacertidae); 2—humerusof Talpa sp.; 3 — jaw of
Desmana sp,; 4,5 — humerus and calcaneus of Lepus зр.; 6, 7 — upper
molars and femur of Steneofiber sp.; 8—jaw of Cricetus sp.
the antiquity ofthe erosion valley and the stability of the hydrological regime
ofthe rivers, which is similar in this respect to the regime of the Don and
the Volga.
The Stavropol complex is not of Upper Pliocene age; Borisyak (1943)
has dated it Pontian. The Lower Pliocene age is indicated by the absence
of elephants and horses proper. According to Argiropulo (1932, 1940c),
the Caucasian Amblycastor is very close to the Upper Miocene species
of this genus occurring in Mongolia and North America. The deer in the
Stavropol faunal complex are of the Lower and Middle Pliocene types. The
Stavropol faunal complex can be tentatively dated as Lower Pliocene on the
bases of accepted geological data and the composition of the complex.
50
(54)
|
Mil 10
= wha
FIGURE 24, Remains of carnivores and proboscideans from the Kosyakin quarry
1,3 — jawsof Ursus cf, arvernensis; 2—jaw of Dinocyon ef. thenardi (x); 4—premolar
of Canidae gen. (x2); 5, 6 — jaw and shoulder of Felis cf. issiodorensis; 7 — Ма Anancus
arvernensis
51
(55)
x
\
= Е
с. |
< ig |
Se Че.
О —=—
FIGURE 25, Remains of hoofed mammals from the Kosyakin quarry
1, 2 —metapodia of Hipparion gracile and Hipparion sp.; 3—jawof Tapirus cf,
arvernensis; 4—jawof Propotamochoerus provincialis; 5,6—horn and jaw of Proca-
preolus зр.; 7—jaw of Cervidae gen,
52
ont
It is younger than either the Taraklian complex of Moldavia (Khomenko,
1914b) or the Pavlodar complex on the Irtysh River and is quite close in
age to the Malusteni complex in Rumania (Simionescu, 1930).
Some geologists correlate the continental clastics, in the foothills of the
Ciscaucasus, 700-800 m thick, with the Cimmerian marine sediments. The
continental clastics in Kabarda and North Ossetia have been described by
FIGURE 26, Jaw of Dicerorhinus orientalis from the Kosyakin quarry
Shvetsov (1928), as follows: the continental sequence resembles a moraine
and consists of loams, sands and tuff breccias with large boulders oferupted
material in some places. The continental beds form a scarp on the northern
slopes of the Chernye-Gory Mountains; in the Zmeika ridge the continental
beds overlie the Sarmatian clays. The beds were probably formed by
mudflows carrying great volumes of poorly sorted material from the
mountain slopes. The fossils in the continental beds are land and freshwater
mollusks (Helix sp., Paludina sp., Neritina sp.), grasses and
leaves, andbones of land vertebrates. M.V. Pavlova has identified an
antler (Cervus (?)) anda "rat jaw'' in Shvetsov's collection.
Sands and clays, 1,000-1,400 m thick, accumulated in eastern
Transcaucasia in Cimmerian time; those on the Apsheron Peninsula
comprise a large oil reservoir, known as the Balakhany productive beds.
There are various opinions on the history of the productive beds.
Kolesnikov, in a review of the theories of their origin (Zhizhchenko,
Kolesnikov and Eberzin, 1940), concludes that the beds were formed ina
"developing tectonic basin, '' whichis $0 зау that they were formed from the
clastic products of the erosion of the eastern Caucasian Mountains which
accumulated in subsiding shallow-water basins. Land vertebrates are very
rare in the productive beds.
In 1932 V. Podgornova discovered fossil mammals in light-colored,
fine-grained sandstones in the productive beds to the east of the Lok-Batan
volcano south of Baku. According to Bogachev's (1938b, 1941) and our
identifications, the following species occur at the locality:
53
58
(56)
Camivora
Vulpes khomenkoi Bog.
Artiodactyla
Cervus (cf. Eucladocerus)sp.
Gazella sp,
The sheatfish (Siluris glanis L.) has also been recorded at the
locality.
Two complete sets of upper molars and jaw fragments of fox have been
found embedded in the light gray, fine-grained sandstone. The bones were
considerably permineralized; the replacing substance is straw-colored or
pink in fresh fracture. The tooth enamel is black and glossy. The material
includes a fragment of gazelle skull with complete sets of upper molars and
two complete metacarpals of deer. There is no basis for identifying, as
Bogachev did, the Middle Pliocene deer from Lok-Batan with the
Quaternary red deer and goitered gazelle. The bones at the locality were
probably deposited in the delta of one of the rivers draining the eastern
slopes of the Caucasus, a river similar to the present-day Sumgait and
Pirsagat rivers.
The layers adjoining the sandstones of the productive beds contain
freshwater mollusks (Golubyatnikov, 1925): Planorbis cornu var.
manteli Dunk., P. costatus Klein, Limnaea armanensis Noul.
and other limnaeids, Unio jasamalicus D. Golub. and Melania
glacilicosta Sandb. The alga, Chara sp., also occurs in these beds.
It is doubtful that such a collection of species could exist under desert
conditions in rivers and lakes of variable salinity. The leguminous plant,
Albizzioxylon hyrcanum, which is characteristic of subtropical
conditions, is known from the Balakhany beds of the Shiraki Steppe
[Leninakan Steppe], west of the Apsheron locality.
The fossils occurring inthe Balakhany beds indicate that the differentiation
of faunal complexes followed the differentiation of the landscapes in the
Caucasus from subtropical forests to hot dry semideserts.
FIGURE 27, Canine (?)of Rhinocerotidae gen, from the Kosyakin quarry
54
The Middle Pliocene mammals of the southwestern part of the Russian
Plain evolved from eastern Mediterranean Miocene faunas. The Middle
Pliocene assemblages of mammals in Moldavia and in the Ukraine are
particularly important to an understanding of the evolution of the Ciscaucasian
fauna. The Moldavian assemblage occurs in gravelly-sandy fanglomerates
in the basins of the Salcia, Kagul and Prut rivers; the Ukrainian
assemblage is known from limestone catacombs, filled with red clay, in
Odessa.
The Middle Pliocene ''fauna'' of Moldaviaconsists of ape, rodent, carnivore,
rhinoceros, Hipparion, hippopotamus, camel, deer and antelope.
Lagomorph and rodent are represented by the Recent genera of hares, pikas,
squirrels, beavers, porcupines and mole rats (Khomenko, 1914a, 1915;
Borisyak, 1943).
. say
AU Tyanovsk iW
te Kuibyshey
ie aS ony
He “tif a
С ae с
Sey a?
, Astrakhan
NY, : Dy
Вашити М4,
= a
FIGURE 28, Kuyal'nitsk and Akchagyl basins (from Kolesnikov; see Zhizhchenko and others, 1940)
Dots indicate main localities
The ''fauna'' of the catacombs in Odessa consists of nearly 30 species of
insectivore, rodent, carnivore, proboscidean, artiodactyl and an ostrich
(Gritsai, 1938, 1939; Borisyak, 1943; Pidoplichko, 1954).
The faunal complexes of the areas north of the Black Sea reflect the
variability of the landforms: wooded creek and river valleys alternating
with steppes. Some of these complexes might have extended southeastward
to Ciscaucasia.
Both the Moldavia (Roussillonian) and the Odessa Middle Pliocene
"faunas'' contain a number of subtropical forms.
The occurrences of ape and hippopotamus in the Roussillonian of the
eastern Mediterranean area are evidence of the warming of the climate in
post-Pontian time. The climate of the north edge of the Cimmerian basin
55
59
60
at the end of the Middle Pliocene was probably similar to the Recent climate
of southern Italy, Spain and France, while in Ciscaucasia the climate was
probably of a more continental type.
During the Upper Pliocene, the configuration of the Caucasian Isthmus
was similar to its present form, but it was intermittently cut off in the north
by narrow straits in the Manych region, which temporarily severed the land
connection between the Caspian Sea and the Black Sea basins.
In the Black Sea region most of the Upper Pliocene localities are situated
along the sea coast. Fossils can probably be found in the zone of contact
between deltaic deposits and gravelly fanglomerates of the paleo-Don, paleo-
Donets, paleo-Dnieper and other ancient streams of the Kuyal'nitsk and
post-Kuyal'nitsk basin (Figure 28).
A number of authors in the last and present centuries (Fischer von
Waldheim (1809), Eichwald (1850), Nordman (1858-60), Pavlova (1895),
Bogachev (1923-24, 1938d), Sherstyukov (1926, 1927) and Grigor (1929)),
have noted the following mammals from the Upper Pliocene of the Azov
Sea and Kuban River regions:
Carnivora Е. meridionalis Nesti
Hypcuas. Е. antiquus Ра]с,
лен Perissodactyla
Rodentia Equus stenonis Cocchi
Trogontherium cuvieri Fisch, Е
Proboscidea Camelus sp.
Cervus sp,
Mastodon arvernensis Croiz. et Job.
Elephas lyrodon Weithofer Bos sp.
Upper Pliocene mammals in the Azov Sea region were collected by
Gromov (1933, 1936) in 1933-1936 in the gravelly sands near the villages
of Morskaya, Merzhanovka and Khapry, and in the Volovaya gulley near
Taganrog. Similar fossils were collected by the author from the Leventsovka
quarries near Rostov in 1954. Fragmentary iron-stained and well-rounded
bones from the Khapry sand indicate that the bones were buried in alluvial
fans and in sediments exposed to marine abrasion. Some of the bones are
marked by bore holes made by marine invertebrates, showing that they had
been deposited in the sea (Figure 29).
According to the identifications of Gromov (1948), Borisyak and Belyaeva
(1948), andthe author, the following species occur in the Azov Sea region:
Proboscidea
Carnivora
Mastodon sp.
Canis cf, lupus L, Elephas planifrons (?)Fale,
Canis sp. E, meridionalis Nesti
Ursus sp.
Hyaena sp. Perissodactyla
Machairodus sp, Hipparion sp
Equus stenonis Nesti(cf. major Boule)
Rhinoceros sp,
Trogontherium cuvieri Fisch. Elasmotherium sp,
Rodentia
56
61
Artiodactyla
Sus sp. Eucladocerus pliotarandoides Alles.
Camelus (Paracamelus)cf, kujalnikensis Cervus cf, elaphus L.
Chom, Cervus sp.
Fragments of ostrich long bones have also been collected in the region
(Struthio sp. ).
| ЕТ i lee
FIGURE 29, Jaw of Elephas meridionalis with borings caused by invertebrates
From the Khapry sands on the right bank of the Don River (Orig. )
Although the species listed are of different ages, as pointed out by
Bogachev (1923-1924), they may, nevertheless, be assumed to represent
one faunal complex. Since the same species also occur farther west, the
complex may be considered characteristic of the Upper Pliocene of the
southern part of the Russian Plain.
Wie eins —IMblosiin IP ey Ta
The burials in the gravels of the Trans-Kuban Plain along the
Psekups River are in the main similar to the northern Azov Sea
localities. The material was collected in 1930-1932 by Gromovy (1935b,
1937, 1948) near the villages of Bakinskaya and Saratovskaya on the left
bank of the Psekups River. Two floodplain terraces can be distinguished
on the bank: the lower terrace of Wurm age andthe upper terrace of Riss age.
Gromov (1948) has demonstrated the presence of freshwater basins in
the foothills of western Caucasia in geological sections which show a
variation from fast-flowing stream deposits to swamp deposits.
According to Gromov (1948, p.56) the following species occur in the
lower and middle parts of the sedimentary sequence:
Proboscidea
Elephas meridionalis Nesti — teeth and limb bones
Dil
62
Perissodactyla
Rhinoceros etruscus Ра]с. — nearly complete skull and skeletal components
Equus stenonis Cocchi — metapodia, phalanges
Equus sp. (aff. stenonis) — limb bones
Artiodactyla
Bison sp. — broken skull
Bos sp. (Leptobos) — broken skull
Eucladocerus pliotarandoides А1ез$. — skull with a horn and horn fragments
Cervus sp, (Rusa) — fragments of horns
Cervus sp. (ex. gr. polycladus?) — fragments of limb bones of a very large deer
Cervus sp. (larger than elk! )
The material also included the lower part of an ostrich tibia (Strutio
sp. ).
The above-mentioned material from the Psekups was lost during the
war before it could be described.
The material preserved in PIN includes an antler of a deer (Eucla-
docerus, Figure 30) and a skull of the Etruscan rhinoceros. The material
is straw-colored and strongly impregnated with iron.
From the presence in the Trans-Kuban Plain of intact skulls and horns,
it can be inferred that during the Upper Pliocene quiet rivers flowed over
this area into large bodies of still water. Such landforms could exist only
at the stage of more advanced peneplanation of western Caucasia.
East of the Psekups there have been individual occurrences of large
Upper Pliocene mammals in the gravel beds deposited in piedmont basins,
such as the bones and teeth of elephant (Elephas cf. planifrons Falc.
and E. meridionalis Nesti) observed along the Laba and Kuban rivers,
in the region of Tshchikskie plavni* (Navozova, 1951), near the villages of
Novo-Labinskaya, Nekrasovskaya, and Grigoropolisskaya (Gromov, 1937),
and south of Armavir. The occurrences of southern elephant, rhinoceros
(Rhinoceros cf. etruscus Jaeg.) and deer (Eucladocerus
pliotarandoides Aless.) are more frequent in the ferruginous sand
near the village of Voskresenskaya, west of Stavropol, andinthe lower part
of the river sands in the Girei quarry near the Kropotkin station (Figure 31).
At the time when this faunal complex lived in Ciscaucasia the climate
was mild-temperate; broadleaf forests of the Colchis and Hyrcania type
covered considerable areas of western Ciscaucasia. Bos, horse and
ostrich fossils evidence a progressive development of savannah-type
grasslands. The animals inhabiting the piedmont plains were probably
often killed by torrential streams and buried in masses of deposited gravel.
In other cases, accumulation of bones in the gravels was caused by the
erosion of bone-bearing lenses by later streams. Therefore, the Psekups
and the Taman faunal complexes (see below) probably antedated the
formation of the bone-bearing gravels in the Ciscaucasian plains.
Most of the fossiliferous strata in the Psekups, Laba and Kuban areas
are usually correlated with the Akchagyl and sometimes with the Apsheron
stage of marine sedimentation in the Caspian region.
* [Periodically flooded areas with Phragmites, Typha and Carex in river deltas and bottomlands in the
southern part of the U.S.S.R.]
58
FIGURE 30. Horn of Eucladocerus pliotarandoides
from the gravels on the Psekups
Taman Peninsula
The bone-bearing formations of the Taman Peninsula are slightly younger
than the lower beds inthe Psekups area. The fossiliferous localities on the
Taman Peninsula are the Sinyaya and Kapustina gulleys and Sennaya, Fonta-
lovskaya and Kuchugury (Figure 32). The bone-bearing breccia at Sinyaya
gulley was discovered by Gubkin (1914) in 1912. He identified the freshwater
sands exposed on the northern coast of the peninsula as of Basal Pleistocene.
The sands contain numerous freshwater mollusks of the genera Vivipara,
Bythinia, Dreissensia, Unio, and other forms (Figure 33). Broken,
sometimes rounded, bones were found to be heavily permineralized,
particularly at the Sennaya and Fontalovskaya localities. Permineralized
excreta of hyenas, swine and ruminants also occur in the sandy-gravelly
beds at Sennaya. The material from the localities mentioned is housed in
59
64
the ZIN and PIN collections, and has been identified by Borisyak (1914),
Belyaeva (1925, 1933a, b), Vereshchagin (1951а, 1957a) and Burchak-
Abramovich (1952a). The species identified are given in Table 4 and
Figure 34.
A vertebra of a large fish was taken from the ferruginous sand near
Sennaya. A seal phalange, resembling those of the Caspian seal (Phoca
caspica Gmel.) (Kirpichnikov, 1953) is probably not from the same beds
from which land vertebrates were taken at Sinyaya gulley.
А taphonomical and ecological analysis of the species studied suggests
that they are part of one faunal complex. The fauna lived on the land which
formed between Ciscaucasia and the Danube River during one of the
regressions in the post-Kuyal'nitsk (Apsheron) basin.
The presence of horse and bison (species adapted to steppe life), of rhinocer-
os, forest antelope, deer andelephant (species adaptedto forest life), andof
beaver (aspecies adaptedto lake andriver life) indicates that western Cis-
caucasia was aplain with dense vegetation alongthe rivers. The climate was
probably moderately warm. From the morphogenetic point of view, this faunal
complex is very close to the Villafranchian complex of Western Europe and
the Nehavend complex of southwestern Asia. However, a number of
FIGURE 31, Chauda and Apsheron basins (from Kolesnikov; see Zhizhchenko and others, 1940)
Dots indicate main localities
Taman species are indistinguishable from the species of the Khapry complex
(beaver Trogontherium, southern elephant, camel) (Vereshchagin,
1957a). Similar species also occur in the Apsheron beds of eastern
Ciscaucasia and Transcaucasia.
The evolution of Upper Pliocene fauna complexes in the Caspian region
was studied at localities situated along the margins of the Akchagyl and
Apsheron basins of the Caspian depression.
In Akchagyl time the axis of subsidence in the northern Caspian became
oriented in a north-south direction, as opposed to its earlier east-west
1704
60
(63) TABLE 4, Upper Pliocene mammals and number of bones from five localities on the Taman Peninsula
Number
of bones
Number
of bones
Carnivora Rhinoceros cf, etruscus Ра]с. 5
Е ff. su b is Wust. 54
Camis памща щен IN ет еее т
Е 305 Wid ois loc о оо © Бека
Е Artiodactyla
Rodentia SUS LAMMAMSMSUS We Were Gos oon0 2
) Camelus cf. kujalnikensis Chom, 1
CASTOR PAMMAMeIMSIS ING Wer osc 46
Г АН Eucladocerus sp. 22
Trogontherium cuvieri Fisch.
Megaceros cf, euryceros Aldr, 1
: Tamanalces caucasicus М. Ver, 3
Proboscidea :
Cervidae реш. 33-55... - - oo00000 41
Elephas meridionalis Nesti* ..... GAZSMA 0 Бо зоовооововоовое 2
В. EROGOMUNEGrAN 199]. оъооросвовв TEAGCIAMNOS GIs ооооооросооос 2
В. AMET@MUS Palle, срообосововоое Strepsicerotini gen, (cf. Taurotragus) 18
Bison cf, schoetensacki Freud. .. 12
Perissodactyla BUGOM Gy». | 0018 0-0 oto во glo BaoloND 2
Elasmotherium caucasicum Boris,
* According to Garutt (1958) this is Phanagoroloxodon mammontoides,
FIGURE 32, Cliffs and landslides on the northern coast of the Taman Peninsula, near Cape Litvinov
Photograph by author, 1954
61
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66
orientation. Akchagyl marine sediments can be traced from eastern
Transcaucasia and the foothills of the Kopet-Dag in the southeast to Kazan
and Ufa in the north (Arkhangel'skii, 1934; Kovalevskii, 1933; Kolesnikov,
(Zhizhchenko, Kolesnikov and Eberzin, 1940)). In Akchagyl time, the rivers
draining the Caucasus Range entered the narrow Kura bay which extended
almost to Tiflis, and the wide Terek bay which extendedto Mozdok. The
rivers transported the bodies of dead animals to the plains and to the
coastal bays.
During the transgressive maximum, the Apsheron and the Kuyal'nitsk
basins were connected by a narrow strait south of the Stavropol Plateau
(Gatuev, 1932).
This growth of the relict marine basin at the time of deposition of the
productive beds has been explained by Kovalevskii (1933) as the result of
marine transgression into the Caspian region. However, his hypothesis
on connections with the boreal sea is not supported by recent data on
molluscan faunas which occur in the northern part of the basin.
Gradual climatic changes probably persisted throughout Akchagyl time.
However, the glaciation ofthe Greater Caucasus, assumed by Kovalevskii
(1936), hadnot yet begun. А reoccurrence of volcanic activity inthe Akchagylis
indicated by thin layers of ash and pumice which appear in the massive
loams and gravels of the foothills region. The Akchagyl basin, which
covered a great area of the Russian Plain, was a natural barrier on the
migration routes between central Asia and Ciscaucasia (Figure 28).
The Akchagyl land flora is known from the shores of the Kura bay, near
Naftalan, from Kakhetia and from the Shiraki Steppe. The following species
have been identified by Palibin (1936): oriential beech (Fagus
Ormemialiis)Moaky (@uwerecus эр. ), willows a lak jalibia)) ‘common
pomegranate (Punica granatum), elm (Zelkova [= Carpinifolia]
crenata), lime (Tilia platyphyllus); alder (Alnus glutinosa),
mulberry tree (Morus andrussovi), Pitsunda pine (Pinus pithyusa),
sequoia (Sequoia langsdorfii) and many other species.
The list of species given above suggests that the environment of the
flora on the shores of the Kura bay was very similar to Recent environments
in the foothills of eastern Transcaucasia.
Palibin (1936) and Grossgeim (1936, 1948) believed that there were no
tropical plants in the Caucasus in Akchagyl time. The last of the evergreen
sequoias and cherry laurels in the Shiraki area probably grew in a humid
climate with moderately cold winters.
The earliest occurrence of Elephas planifrons and Equus
stenonis inthe Caucasus is in Akchagyl beds, although land vertebrate
fossils seldom appear in these beds and the material mainly consists of
teeth of mastodon, elephant and horse.
In Ciscaucasia, teeth of Mastodon arvernensis Croiz. et Job.
and Elephas planifrons Falc. were collected on the Malgobek ridge,
40 km south of Mozdok. Teeth and bones of Elephas planifrons (?)
were also found in the Solenyi gulley near Grozny (Pavlova, 1931).
In eastern Transcaucasia, the Akchagyl conglomerates between the
Alazan and Kura rivers (Kudryavtsev, 1932) contain bones of large
vertebrates, freshwater mollusks (Helix sp., Planorbis,
Cyclostoma) and species of plants known in the Recent. An antler of
Cervus sp. is known from the Akchagyl of western Azerbaidzhan.
The coarse breccia of the Bedeni ridge in the Tsalka District of Georgia
contains fragmented bones of mammals among which teeth of Elephas aff.
63
FIGURE 34, Fossils of the Taman complex
1 — дам of Canis tamanensis; 2 — ulna of Panthera sp.; 3—skull of Castor tamanensis;
4 — иррег jaw of Trogontherium cuvieri; 5 —upper jaw and teeth of Elephas meridionalis;
6, 7 —Pmgand metacarpus of Equus aff.stissenbornensis; 8—jawof Sus tamanensis; 9—horn
of Eucladocerus зр.; 10—horn peduncle of Tamanalces зр.; 11 — Пот axis of Bison sp.
64
68
planifrons Fale. and Equus stenonis Cocchi have been identified.
The breccia is underlain by a layer of doleritic lava and is covered by
lacustrine sands and clays. The lake sediments are overlain by a layer
of dolorite (Zaridze and Tatrishvili, 1948). Thus, in that area, the mammals
lived and died during a period when the volcanos in the Lesser Caucasus
were dormant.
The next transgression in the Caspian
=—_ м Basin, a somewhat smaller one, is known
as the Apsheron sea.
The Kura bay of the Apsheron sea
08 reached the longitude of Kirovabad. The
Terek bay was temporarily connected with
the Euxinic basin by a strait in the Manych
area. The sea reached the latitude of
145 Sarepta and [Lake] Inder in the north.
2 The climate and landforms of the
Caucasus in Apsheron time probably
Е бо remained the same as in the Akchagyl, and
а: Oe >. the volcanic activity was of about the same
Ни: : intensity. Torrential mudflows, caused by
| heavy rains, carried volumes of gravel
and boulders from the mountains
(Kudryavtsev, 1933); these boulders can
now be seen on the Kakhetia Plain.
The land vegetation known from the
Apsheron deposits in the Shiraki Steppe
consisted of spruce (Picea orientalis)
and a number of Recent forms: beech,
oak, aspen, apple, willow, filbert, Turkish
filbert, walnut (Juglans regia),
zelkova, honeysuckle; and Hyrcanian forms:
oak (Quercus castaneifolia), alder
(Alnus subcordata), maple (Acer
ibericum) (Palibin, 1936). Cooling of the
climate in Apsheron time resulted in the
disappearance of the subtropicalevergreens.
The wooded landscape of the shores of
the Kura bay was probably similar to those
parts of the present-day landscapes of
Colchis and Asterabad which are stillina
completely natural state.
During the regressions of the Akchagyl
and Apsheron seas, arid conditions
undoubtedly prevailed in the eastern part
of the Caucasian landmass. The
development of coastal steppes and
semideserts created the environment for
ferruginous sands with rounded fragments us pei Sail ou Sireigiger Sine) ФА, weighs.
of volcanic ash, bones of elephants and Fossil mammals are more abundant
hoofed mammals, Numbers on the right in the Apsheron than in the Akchagyl
indicate thickness in meters sediments. The material consists mostly
a meth Pin Е Cie
5 =e СВ RCS Os
Е
а
м РЕ...
‘о’. .’о’..0.- ‘о. *.`Р. 2 ct
Pere gor, Soy Ao eo : 2 One
=
о
о.
[©
FIGURE 35. Stratigraphic section of
bone-bearing sands near Georgievsk
a-c — gravels with obsidian fragments on
the high terrace; d-i — light-colored and
65
69
of single limb bones and teeth of proboscideans and perissodactyls. These
burials occur primarily in littoral marine deposits formed in quiet bays
at various basin levels. Some occurrences are known from continental
deposits: from the zone of accumulation at the piedmont ''barrier'' at a
mountain base, and from the travertines in the Pyatigor'e area.
In eastern Ciscaucasia, ''traces'' of a faunal complex, similar to those
of the Psekups and Taman, occur in the travertines and loams of Mount
Mashuk, and in the coastal loams of the Apsheron basin. Bones of elephant
and hoofed mammals from the travertines are brittle and some contain
cavities formed by dissolution, and filled with calcite crystals. From
the oldest, Upper Pliocene ''travertine complex'’ in Mount Mashuk, the
following forms have been described: Elephas meridionalis Nesti,
Е. cho antiquus) Fale, Ey aff: tregont her Wust, Bovinae, Cervinae
(Egorov, 1932; Ivanova 1948; Gromov, 1948).
We collected teeth of Mastodon arvernensis Croiz. et Job.,
southern elephant (Elephas meridionalis Nesti), horse (Equus
ef. stenonis) and bones of deer (Eucladocerus sp.) (Figure 36)
from alluvial gray sands below a layer of redeposited basalt fragments and
volcanic ash (Figure 35), near the Cossack village of Vinogradnaya and
near Georgievsk.
Bones of Elephas meridionalis and Equus sp. have been found
in the Apsheron loams on the slope of Mount Tash-Kala near Grozny
(Pavlova, 1931). Twoteeth of Equus stenonis Cocchi and an antler
of a large antelope (Bubalis sp.) found near Verkhni Achaluki on the
Sunzha ridge are probably also of Apsheron age (Burchak-Abramovich,
1952a).
The following forms from the locality near the village of Vinodel'noe,
northeast of Stavropol, are identified by Borisyak and Belyaeva (1948) in
the PIN collections:
Proboscidea
Elephas meridionalis Nesti
Perissodactyla
Equus (stenonis?)sp.
Artiodactyla
Paracamelus gigas Schloss.
A camel mandible from this locality has been described by Khaveson
(1954b, Figure 37).
Teeth of southern elephant from the piedmont gravels near Makhachkala
have been recorded occasionally.
Apsheron Peninsula
Fossil marine and land mammals are very abundant in the limestones
of the Apsheron stage on the Apsheron Peninsula near Binagady, Shikhovo
and Baku.
66
М. от у
И 7
vt HH
| if ' qj
[0
er
Он ПИН.
Ч
т: НЕХ \
АИТ
Ц
и,
— Al > nl |
Ще i lim
И
FIGURE 36, Fossils from the river sands in the Georgievsk quarry
i 2—M? of Elephas meridionalis and worn surface of tooth; 3, 4— worn surface and inner
surface of Pmgof Equus stenonis; 5—metacarpus of Eucladocerus sp.
67
Г
According to Bogachev (1923, 1938a, 1940d), the fauna includes
porpoise (Delphinus delphis L.), an unusual predacious marine
pinniped similar to otter and seal (Necromites nestoris Bog. ) and
perissodactyls (Hipparion crassum Gerv., Hipparion sp.). The
humerus of a seal, indistinguishable from that of Phoca caspica Gmel.,
has been found under a 22-m-thick bed of Apsheron limestone near
Baladzhar. The presence of BlackSea porpoise (Delphinus delphis L.)
in the Caspian region can be accounted for (Bogachev, 1939a) by migration
from the Mediterranean in Akchagyl time, i.e., at the time of migration
of the mollusks, Mactra, Cardium and Potamides. According to
Bogachev, the dolphin became extinct in the Caspian at the end of the
Apsheron because of advancing glaciation.
The same can be said of the incompletely studied Necromites which,
according to Bogachev (1940d), is similar to Semantor macrurus
Orl. from the Pliocene deposits of eastern Siberia.
Farther west, fossils occur in the clayey ridges of Kabristan and in the
two or three parallel ridges of hills which extend from the lora and
Kartalinia plateaus to the Gerdyman-Chai canyon in the east. The hills
are made of clays and loams with gravelly intercalations totalling 400 m
in thickness.
The material collected by geologists in the Kondzhashen, Palan-Tikyan,
Karadzha, Boz-Dag, Chuzgun-Tapa and other ridges has been identified
by Bogachev (1938c), Burchak-Abramovich (1951а, 1952c), and the author.
Rodentia Perissodactyla
Trogontherium cuvieri Fisch, Equus sp,
Muridae gen. Rhinoceros cf, etruscus Ра]с.
Rhinoceros sp,
Proboscidea
Artiodactyla
Anancus arvernensis Croiz, et Job,
ane Е - Cervus :
Elephas ineridionalis Nesti ay
Eucladocerus sp,
Bison sp,
FIGURE 37, Jawof Paracamelus gigas from Vinodel'noe
68
74
It is possible that the upper jaw of Ste godon sp. in the collections
of the Natural History Museum in Baku came from some Upper Pliocene
deposits in eastern Transcaucasia (Bogachev, 1935).
The most abundant Upper Pliocene species from the Apsheron beds of
Transcaucasia are, as a whole, very similar to the post-Kuyal'nitsk fauna
of eastern Ciscaucasia. The mastodons occurring in the Apsheron beds
were either the last survivors of the eastern Mediterranean group, or they
were redeposited from older beds.
Upper Pliocene index mammals have been identified from near Leninakan
and Erzurum in the Armenian-Anatolian highlands of southern Transcaucasia.
The fossils probably occur in the gravelly-pebbly sands deposited by streams
in small, inland, freshwater lakes.
As early as 1899, Abich mentioned an accumulation of fossil bones in Upper
Tertiary calcareous marls and conglomerates underlying the layer of tuff
and lava onthe slopes of a fortified hill near Leninakan. Academician
I. Е. Brandt has identified ''Mastodontidae, Cervidae and Bovidae'' in that
material.
Bogachev (1923-1924) mentioned the finds of Elephas armeniacus
Fale. in gray volcanic sands near Leninakan. The species is probably
identical with ЕВ. meridionalis Nesti.
In later collections from that area Burchak-Abramovich identified bones
of Elephas sp., Rhinoceros mercki Jaeger, and Equus
stenonis Cocchi (Avakyan, 1948).
The following list (which requires revision) is given by Bogachev (1938c)
for species occurring in the Upper Pliocene diatomite near the village of
Nurnus on the Zanga River:
Carnivora Rodentia
Mustela filholi Gaudry Gerbillus sp.
Putorius sp.
Perissodactyla
Lagomorpha Hipparion sp.
Lepus sp. Rhinoceros etruscus Falc,
Artiodactyla
Tragocerus sp.
The diatomite, 7m thick, occurs between andesitic lavas below and basalt
above (Gambaryan, 1934), indicating that a freshwater basin existed in the
middle Zanga region in a period between two strong volcanic eruptions.
Intercalations of pumice and ash in the diatomite bed show that the volcanic
activity also continued during the deposition of the bed.
Occurrence of gerbil bones indicates that xerophilous vegetation thrived
along the southern margins of the Armenian Highland in Upper Pliocene time.
The stratigraphy of the fossil localities and the species composition of the
Upper Pleistocene Transcaucasian fauna indicate that in Apsheron time the
climate was temperate, and the volcanos were very active. These data also
reflect to a certain degree semi-arid to arid conditions in the southern
and eastern parts of the country.
Pleistocene elements are traceable in the Upper Pliocene faunal complex
of the Caucasus, particularly in the Taman Peninsula.
69
(72)
TABLE 5. Stratigraphic distribution of the main localities of Tertiary mammals in the Caucasus
Division Localities: Black Sea Area and Ciscaucasia
Chauda beds Taman Peninsula: Sinyaya gulley, Sennaya,
Kuchugury
Gurie beds
Kuyal'nitsk
=
Meotian
Stavropol area: Voskresenskaya, Trans-Kuban
Plain: Psekups, Bakinskaya
Taman Peninsula: Kapustina gulley
Pliocene
Zmeika ridge; Darg-Kokh
Stavropol Plateau; Kosyakin quarry
North Azov coast: Novocherkassk, Tuzlovka
Sarmatian Stavropol area; Burlatskoe, Blagodarnoe,
у Petrovskoe
Miocene
Konka
Karagan
Chokrak
Kuchugury
Helvetian
Stavropol area: Belomechetskaya
Oligocene
70
Localities: Caspian Area and
Transcaucasia
(73)
Stage
Akchagyl
Productive beds
Pontian
Meotian
Terek Range: Malgobek, Grozny,
Achaluki
Apsheron Peninsula: Binagady, Khurdalan,
Shikhovo
Armenian Highland; Leninakan, Nurnus
Тога Plateau: El'dar, Dzheiran-Cheli
Kirovabad Plateau: Karasakhkal, Naftalan
Iora Plateau: Palan-Tikyan
Apsheron Peninsula: Lok-Batan
Kartalinia Plateau: Dzhaparidze
Gori valley: Arkneti
Iora plateau: El'dar, Udabno
Kartalinia Plateau; Kyasaman,
Kotsakhuris- Kedy
Sarmatian
Konka
Karagan
Chokrak
Helvetian
Predominant Groups of Species
Taman wolf, southern elephant,
Trogontherium, Equus
stenonis, Cervus pliotaran-
doides, Taman bison
Mastadon arvernensis
Aceratherium
Hipparion
Cervus pliotarandoides
Fox,
deer,
gazelle
Dinocyon,
Mastodon arvernensis,
hamster, Dicerorhinus,
tapir, Hipparion, roe deer
Hyena, long-snouted mastodon,
Aceratherium, Hipparion,
giraffe, gazelle
Amphicyon, swamp mastodon,
Paranchitherium,
Anchitherium, giant swine,
antelope
Dzhavakhetia: Akhaltsykh
71
Benaratherium
Chalicotherium
Anthracotherium
Tragulids
75
The Neogene faunal complexes of the Caucasus are not endemic. The
Oligocene ''fauna'' of the Lesser Caucasus is related to the central Asian
and European ''faunas'' of the same period.
The Middle Miocene "fauna" of Ciscaucasia, known from Belomechetskaya,
is also related to the Miocene ''faunas' of Eurasia and North America. This
"fauna'' migrated over temporary land bridges from the north and south,
although the trend from the south was more pronounced.
In the Upper Miocene, Sarmatian, the Hipparion complex migrated
to the Caucasus from the south over land which joined the island of
Caucasus with Asia Minor.
The local fauna of the mountains has probably evolved since the Miocene.
Some of the surviving, endemic elements of the highland fauna are
undoubtedly descendants of Upper Miocene forms.
Evolution of the Pliocene faunal complexes in the piedmont and the
plateaus can be traced mainly to migrations from the north and south.
The hot, dry climates of the Pliocene produced a more pronounced southern
influence on the evolving fauna.
It is only at the very end of the Pliocene that the northern faunal elements
became important. So-called Pleistocene genera and species appear during
the general cooling at the end of the Pliocene. Evolution of endemic species
in the Pliocene occurred mainly in the highland regions and left relicts on
the peneplain.
The paleontology of the Caucasian land mammals does not support the
hypothesis of a ''Pontian landmass" in the Black Sea Basin connecting
western Caucasia, the Crimea and Asia Minor (Andrusov, 1918; Vul'f,
1944; Puzanov, 1949). This hypothesis has also been discarded by geologists
(Muratov, 1951). However, it is more than likely that dry land existed in
the Upper Pliocene where the Sea of Azov now lies (Vereshchagin, 1957).
A stratigraphic summary of Caucasian localities of Tertiary land
mammals is given in Table 5.
Many forms of the Upper Pliocene Taman complex survived in the
Caucasus and continued to evolve through the Lower and Middle Pleistocene.
72
76 Chapter II
77
DEVELOPMENT OF CAUCASIAN LANDSCAPES AND
MAMMALIAN FAUNA IN THE QUATERNARY
EE ee ОО bN Ee РН st OC 2 NE ws On Diy
АЛЕ ЕАО СА Suis
The specific features of evolving landforms in this area between two
ancient continents make the identification of the Pliocene- Pleistocene
boundary difficult.
The variety of zoogeographic zones and the complexity of their boundaries
developed in the Upper Pliocene as well as in the Recent.
Therefore, any study of the geomorphology based upon single occurrences
of plants and animals in continental deposits often reflects only local
conditions.
This problem, as it concerns the European continent, is still under
discussion by geologists and biologists (Zhirmunskii, 1936; Nikolaev, 1947,
1950). We agree with Nikolaev that the Pliocene-Pleistocene boundary must
be identified on the basis of all available data of the various disciplines,
rather than on isolated criteria.
A variety of geomorphological changes caused by the interplay of
tectonics and climate have been proposed as the markers of the Pliocene-
Pleistocene boundary. Such markers are sea level fluctuations, orogenic
and erosional cycles, glaciations, evolutionary changes of plants and
animals, and morphogenetic stages of fossil lineages.
The lower boundary of the Quaternary has been variously placed — in the
Upper Sarmatian by geologist Kovalevskii (1936) and botanist Grossgeim
(1936); at the end of the Akchagyl by geologists Reingard (1931, 1936a),
Gerasimov and Markov (1939) and paleontologist and geologist Gromov
(1948); and in the Upper Apsheron by geologists Pavlov (1936), Mirchink
(1936b) and Vardanyants (1948).
The Russian zoogeographers Menzbir (1934) and Serebrovskii (1935)
did not resolve the question when they applied Western European
stratigraphic concepts to the Caucasus.
The hypothesis of Kovalevskii, Grossgeim and Shvetsov (1928) that the
Pliocene continental glaciations in the Caucasus correlate with the Gunz
and Mindel glaciations is no longer accepted.
Gromov (1948) inferred fromthe geomorphological and paleontological
data that, sometime atthe Tertiary-Pleistocene boundary, the climate
became more severe which, however, did not imply an extensive glaciation.
He considered the Khapry and the Psekups faunal complexes (see Chapter I)
characteristic of the transition from the Pliocene to the Pleistocene in
73
78
Ciscaucasia and the southern part of the Russian Plain, and placed the
Taman complex at the base of the Quaternary. *
At that time the following forms were considered indicative of early
preglacial time on the Caucasian Isthmus: Elephas trogontherii
(Е. wisti М. Pawl.), Bison schoetensacki Freud., Elasmo-
therium caucasteium Boris. ,Cricetus остова i, Spokes
microphthalmus Guld. The faunal break was expressed in the
disappearance of mastodon, elephant (Elephas planifrons and
E. meridionalis), Hipparion and Etruscan rhinoceros, coupled with
the appearance of bison. Thus Gromov's biostratigraphic criteria of the upper
margin of the Tertiary are similar to those of West European authors: the
Khapry and Psekups complexes coinciding with the Gunz and Gunz-Mindel
glaciations, and the Taman complex with the Mindel glaciation of the West
European glaciological scheme.
The first erosional cycle in the Quaternary occurred in the Gunz. The
cycle is represented by gravels of the Kuban River terrace which are
175° mifhick,
Similar paleontological considerations have been put forward by
Pidoplichko (1940c, 1952, 1954) for the northern Black Sea coast and the
Ukraine.
New paleontological data have confirmed that pronounced qualitative
changes in the flora and fauna of the Caucasian Isthmus occurred only at
the end of Apsheron time. These changes are indications of a mildly warm
or temperate climate.
Silicified wood of broadleaf species — oak (Quercus sp.), beech
Pliocene freshwater beds near Kuchugur on the Taman Peninsula, and in the
lower part of the alluvial sands in the Girei quarry on the Kuban. It is
quite possible that subtropical species did not exist in the Caucasian flora
of this period. An abundance of terrestrial mollusks (Chondrula,
Helix) and freshwater mollusks (Corbicula, Anodonta, Bithynia,
Micromelania and other species) in the post-Kuyal'nitsk continental
beds of the Taman Peninsula and in the Upper Apsheron beds of eastern
Transcaucasia, is evidence ofa temperate climate.
It was in Upper Apsheron time that mastodon and Hipparion became
extinct in the foothills of the northern Caucasus. The forms which then
appeared were horse (Equus cf. stenonis, Е. aff. sussenbornen-
sis), Elasmotherium caucasicum, camel (Paracamelus)
and bison (Bison sp.).
The morphogenesis of the Caucasian elephant (Elephas meridiona-
lis —E. trogontherii) was in a transition stage during the Upper
Apsheron (Vereshchagin, 1957a).
The picture is similar in the continental deposits of the Transcaucasus,
particularly so in the Armenian Highland. However, the chronological
correlation with the Ciscaucasian localities is uncertain, despite the
similarities in the composition of the faunal complexes and in the
morphogenetic stages.
* When Gromov wrote this, the Taman complex was poorly known; in point of fact, it is very similar to
the Psekups and Khapry complexes (Vereshchagin, 1957a),
** ‘Identified by А.Г. Zubkov.
74
79
Thus at the end of Apsheron time, new cycles of erosion and mountain
formation commenced on the Caucasian Isthmus; the flora developed a
temperate or, in some places, a xerophilous aspect, and the vast areas
from which the seas receded came to be inhabited by ungulates of a type
adapted to the steppe, or even to the desert.
Taking into account that this continental phase undoubtedly covered a
very long period, we agree with Pavlov (1936) in placing the beginning
of the Anthropogene in the Caucasus at the end of the Apsheron sea
regression and at the beginning of the Baku sea transgression.
THE PROBLEM OF THE CAUCASIAN CLIMATES AND
LANDSCAPES IN THE PLEISTOCENE AND THEIR BEARING
ON THE EVOLUTION OF FAUNAL COMPLEXES
The chief considerations in studying the development of the climates and
landforms of the Caucasus in post-Apsheron time are the following:
1. Tectonic cycles and the dynamics of the land relief created by river
erosion.
2. Fluctuations of the sea level and the land connections between the
Ciscaucasus and the Russian platform.
3. The extent and number of glaciations as related to the tectonics and
climatic fluctuations.
4. Changes in flora and formation of vegetation zones.
These factors have been studied in the Caucasus on an empirical basis,
disregarding the paleontological material. A summary of these studies
would be germane to the overall problem of the history of the fauna.
Dynamics of land relief. Inthe Pleistocene the dynamics of the
relief of the Caucasus was highly complicated. The peneplanation of the
ancient mountains of the Greater and Lesser Caucasus at the end of the
orogenic period of the Top Pliocene (Apsheron age) is a fact accepted
by geomorphologists.
Remains of an ancient peneplain are in fact observable in the Main
Range, e.g., in Abkhazia, Svanetia and Kabarda, and particularly in
Dagestan. At the end of the Pliocene, the poorly drained, boggy alpine
plateaus of the present Dagestan highlands were probably no higher than
500-800 m above sea level. The peneplain is considered to be of either
Apsheron or Baku age; Mirchink (1936a) regards it as Upper Apsheron-
Lower Baku. Some ridges in the region of Vodorazdel'nyi and the Bokovye
ridges presumably did not rise over 1,000-1,500 m above the level of the
Apsheron sea. Plant and animal fossils from these plateaus would have been
of great interest in the study of the faunal complexes. Unfortunately, their
preservation was probably precluded by the action of rapid drift and erosion.
According to Nikolaev (1941, 1949) the relief of the inner plateaus of
the Greater Caucasus is much older than Apsheron time. The relief of the
Armenian Highland (Paffengol'ts, 1948, 1950) can be traced as far as the
time in the Lower Quaternary when Lake Sevan was formed by the lava
flows from the Bogu-Dag volcano which dammed the deep erosional valley
of the paleo-Zanga River.
Nikolaev (1941, 1949) places the maximum uplift of the Greater
Caucasus, following a series of oscillating, tectonic movements, at 3-4 km,
75
80
which was reached in ''post-Upper РПосепе"' time. On the basis of their
studies in Dagestan and Ossetia, Reingard (1939) and Vardanyants (1933b,
1948) hypothesized that the central part of the Main Range is a horst,
which was uplifted by 1,000-1,500 m. The figure of 3-4 km is probably
the sum of the uplift of the Central Range and the subsidence in the Black
Sea and Caspian Sea depressions.
The most recent tectonic movements have been detected in the central
Caucasus, in the Stavropol area and on the Taman Peninsula. The laccoliths
in the Pyatigor'e area, which stand out in the surrounding relief, ''underwent
a relatively greater uplift than the surrounding areas.'' Ronov (1949),
working by a ''volume method,’ stated that the uplift of the Greater Caucasus
in the Pleistocene was nearly 2,000 m.
Thus, by a general consensus, the relief of the Caucasian mountains
resulted from arching, or, in some places, from horst uplift, and from a
concomitant subsidence in the piedmont plains and in the Black Sea and
Caspian Sea depressions.
Because of these tectonic movements, the younger terraces in the
foothills were formed close by older, denudational terraces, which, in the
plains, disappeared completely under the younger ones. This overlapping
of terraces is very important in a stratigraphic correlation of terrace
deposits and in fossil dating.
The uplift of the Caucasian mountains inevitably resulted in glaciations
(which are discussed in more detail below) and in the formation of a rocky
relief with numerous cuestas and talus in the faulted areas. The new
topography allowed an expansion of plant and animal species adapted to a
rocky terrain. The higher relief also promoted a greater ecological and
morphological differentiation which was reflected in the formation of new
habitats and in the division of existing ones.
The slopes of the ridges which rose during the Pleistocene were actively
eroded by rivers. Mountain streams deepened their valleys, stage after
stage, forming terraces which correspond to the mountain-building cycles.
Rivers cut through the young rising ridges forming narrow canyons in the
process.
Such post-Pliocene canyons occur on the northern slope of the Main
Range (along the Baksan, Chegem, Cherek and Urukh rivers); in eastern
Transcaucasia (along the Geok-Chai, Tur'yan-Chai, Alazan, Kura and
Mtskheti rivers); on the southeastern spurs of the Karabakh Mountains
(along the Araks River), and in Talysh (along the Vazaru-Chai river).
The differential uplift of the mountain ridges combined with river erosion
produced sheltered valleys, screened by ridges, which captured atmospheric
precipitation and resulted in the development of temperate, arid areas in
the highlands. These were a refuge for plants and animals adapted to
drought. During the Pleistocene such areas existed in Dagestan, in the
longitudinal mountain valleys of Ossetia and Kabarda, in the Gori depression
on the Kura River, in the upper part of the Pambak River valley on the
Armenian Highland, andin the Zuvanda depression in Talysh, These areas
are still inhabited by xerophilous animals: suslik, hamster and steppe vole.
Karstic niches and caves which were created in limestone terrain by
river erosion became habitations for some predator species and, later,
for man.
A thick mantle of gravels and loams formed in the piedmont during the
Pleistocene from the transport of clastic material by rivers from the
76
81
82
mountains. Such deposits cover the Trans-Kuban and Terek-Sunzha valleys,
the Kusary and Kirovabad plains in Transcaucasia, and the terraces on the
Black and Caspian seacoasts.
These piedmont plains were inhabited by mammals associated with the
forest-steppe and with moderately humid climatic conditions. The piedmont
plains were also the paths of migration from north to south, and vice versa,
of those species which lived on the lowland plains.
Vulcanism has had a pronounced effect on the relief and hence on the
areas of species distribution in the highlands and on the piedmont plains
(Apsheron and Taman peninsulas). In the Armenian Highland the lavas
covered great areas of the plateau, sometimes nearly 50 % of awhole area
(Maruashvili, 1946). Volcanism affected the distribution of animal species
both directly and indirectly. For example, the absence of forests inwestern
Armenia is, according to some geobotanists, the result of the volcanic
activity and lava flows of the Pleistocene (P. Yaroshenko, 1941).
The orogenic cycles were probably closely related to fluctuations in
sea level during the Quaternary.
Fluctuations of sea level. The transgression of the Baku sea,
which followed the Apsheron regression, was characterized by an abundance
of Didacna. The next stage, the Khazar sea, was characterized by the
archaic Didacna surachanica Andr. The equivalent in the Black Sea
Basin was the Neo-Euxine sea. The Khazar sea was followed by the
brackish Khvalynsk sea which extended far to the north leaving brackish-
water mollusk fauna near Ural'sk and Kazan. The Khvalynsk sediments
correlate with the beds of the Ancient Black Sea. The last major
transgression in the Caspian depression (Khvalynsk sea) is usually thought
to correspond to the last stage of glaciation of the Caucasian Range. The
Khvalynsk sea receded into the postglacial Caspian, in which later trans-
gressions were only minor and did not extend much beyond the present
coastline (Kovalevskii, 1933; Figure 38).
The major Pleistocene transgressions considerably reduced the coastal
terraces on the east and west coasts of the Isthmus.
Sea erosion of the piedmont barrier, as in eastern Transcaucasia, or
of the coastal plains restricted the areas of distribution and migration of
animals on the plains. The alluvial plains were also affected by marine
erosion: the Yeisk-Kuban, Kuma-Terek, Rion and Kura-Araks plains,
which, at various times, were inhabited by faunas of diverse origins.
The Manych strait formed during the transgression must have restricted
the migration of animals from the Russian plains at least to the extent that
hibernating animals could not have migrated over ice-covered country.
During the regressions the low-lying areas remained covered by broad
salinas and marshes with residual lagoons and lakes. Desalinization of
these areas progressed rapidly in western Ciscaucasia where steppes
developed. In eastern Ciscaucasia, desalinization proceeded at a slower
rate. There semideserts developed, inhabited by jerboas, gerbils, saigas
and goitered gazelles. Residual lakes and lagoons became plavni overgrown
with nymphaea and bulrushes. Colchis developed a marsh-type vegetation
consisting of alder, nymphaea, sedge and sphagnum. The longevity of such
marshy tracts depended on the hydrological regime of the feeding rivers
and the rate of alluvial fill. The fauna of these marshes included such
hydrophilous animals as otter, water rat, European beaver and boar.
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FIGURE 38, The Caucasus and the Caspian in the Upper Pleistocene
1 — old freshwater sediments in the Volga Basin; 2 — Aral-Caspian marine
transgression; 3 — region of Quaternary glaciation (from Kovalevskii, 1933)
Extent and number of glaciations. Climatic fluctuations and
glaciations in the Caucasus could have been the most important factors in
the development of the fauna.
Data on Caucasian glaciations is based on the geomorphology of the area
and on the relationships between the glacial and orogenic cycles and the
sea level stands. Correlation of these data has been complicated by the
attempts of Russian geologists to identify Caucasian glaciations with the
European glacial stratigraphic scheme developed for the Alps. There has
been almost no original stratigraphic work done to date on the glaciations
in the Caucasus that is free of this ''Western glaciological hypnosis."
Geomorphological studies show those peculiarities which distinguish the
development of the Caucasus during the Pleistocene from the development
of the Alps.
Academician Pavlov (1925) believed that the end of Apsheron time in the
Caucasus coincided with the Pliocene (Gtinz) glaciation, and that it was marked
by the uplift of ranges causing a formation of mountain glaciers and intensive
deposition of the gravels which are common in the upper part of the
Apsheron suite. According to Pavlov, the Mindel glaciation correlates
with the upper Baku and the Riss glaciation with the Upper Khazar, while the
78
83
84
Khvalynsk corresponds to the time when the Wurm glaciers waned.
Essentially this correlation is still valid.
Reingard (1937, 1947b) recognized four or five major stages of glaciation
in the Caucasus. The first (Gunz), which glaciated as far as the piedmont,
occurred in the transition from Akchagyl to Apsheron time, and is
represented only by scattered gravel deposits. The Mindel glaciers extended
80 km north on the piedmont plain of the central Caucasus during the
Apsheron-Baku transition period. This glaciation was followed by the long
second interglacial, during which thick sequences of diluvial loams
accumulated from transport of clastic material from the mountains. The
third glaciation, the Riss, was the most extensive and its erosional effects
in the valleys were the most pronounced. During the interglacial which
followed, the glaciers remained only on the mountain peaks. During the
fourth and the last glaciation, the glaciers covered only the mountain
valleys, and subsequently receded in four stages, with one advance which
occurred in the Middle Ages of our era.
According to Reingard, the rise in the level of the Caspian Sea was
related to the decrease in evaporation at the time of glaciation. Initially,
the sea level rose slowly. Later, during the melting of the glaciers, the
rise was more rapid, attaining its maximum in the second half of the glacial
phase. At that time, the Black Sea level had not yet reached its maximum
(Figure 39).
According to Gerasimov and Markov (1939), the entire glacial history
of the Caucasus can be subdivided into two periods. During the first period
(Akchagyl-Apsheron), one or two glacial phases occurred in central and
eastern Caucasia as the result of the uplift of the mountains. In the second
period (the second half of the Quaternary), two glacial phases developed in
the Greater Caucasus, ''the mountain framework formed by the Lower
Quaternary tectonic movements. '' The authors doubted any synchrony
between these and the alpine glaciations, although they were certain of
"basic changes in climate which occurred several times, along witha
number of glaciation (3-4) in the Caucasus. "
A record number of ten glaciations beginning with the Upper Miocene
has been given by Kovalevskii (1936).
Three or four glacial phases in the Caucasus have been suggested by
Vardanyants (1948). However, he tended to accept Gromov's (1935b)
subdivision into two glacial epochs corresponding to the Riss and Wurm of
Europe: the Riss corresponding tothe Early Khazar, andthe Wurm tothe Early
Khvalynsk, with corresponding stages of retreat following each (Figure 40).
In the Black Sea Basin the periodically established connections with the
ocean were controlled by the uplift and subsidence in the Bosphorus -
Dardanelles region and modified the processes of glaciation and
transgression. A stratigraphic summary of Vardanyants' is given in
Table 6. *
A number of marine basins periodically connected the Aral, Caspian,
Black and Mediterranean seas, and their waters interchanged with the
fresh waters of the Caspian through the Manych strait. Increase in salinity
of the basins was due to the penetration of the Mediterranean water eastward
in post-Tertiary (Karangat and Ancient Black Sea) time.
* Other geologists, for example Popov (1955), correlate the Khvalynsk time with the Neo-Euxine beds,
the Khazar time with the Karangat beds, etc,
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86
Ц seems strange that the views of geologists оп {Ве extensive glaciations
in the Caucasus were supported, rather than opposed, by biologists.
CASPIAN
:
CAUCASIAN TRANSGRESSIONS
GLACIAL AND INTERGLACIAL rae
STAGES REGRESSIONS
; é Aral-
Cardium edule basbiekiibelt
FF [amr nase
В | Stages of retreat =| Didacna trigonoides | <span
beds
Wirm glaciation ——_ = =
Riss- Wurm- D surachanica Middle
interglacial Caspian
. 2 beds
Kalinovka orogenic
phase
D. carditoides
Pre-Riss orogenic
phase
MR Long Mindel-Riss ==
glaciation 5
Baku beds
Post-Gunz orogenic
Mindel glaciation phase
Gunz-Mindel
interglacial Upper
Vs Apsheron
Gunz glaciation
Preglacial time — Lower Apsheron
FIGURE 39, Relationships between glaciations, orogenic phases in the Caucasus and fluctuations
of the Caspian Sea level (according to Reingard, cited in Bogachev, 1940)
Formation of vegetation zones. According to the botanists
Kuznetsov (1909) and Medvedev (1915) and the zoologists Satunin (1910)
and Dinnik (1911), there was only one glacial phase during which the
Caucasian fauna and flora were saturated with northern forms, and the
mountain animals and plants had to seek refuge on the plains.
From an analysis of Recent Caucasian flora, Grossgeim (1948) has
inferred a number of hot interglacials, in order to account for the presence
of xerophilous southern plants at a number of places in the Caucasus.
Geobotanists Maleev (1946) and Kolakovskii (1947b), who studied the origin
of the eastern Mediterranean flora (in particular, that of Colchis), also
80
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Я
Novorossiisk
о
Stavropol
di
Elbrus
Makhachkala
«>
Tbilisi
о
Batumi
—5 adil
2
Yerevan ©
FIGURE 40, Glaciation in the Caucasus during the pre-Khazar orogenic phase
1 — inferred outline of the sea; 2 — areas of glaciation (from Vardanyants, 1948)
accepted the notion of several glacial epochs and warm interglacials. They
connected the occurrence of mesophilous broadleaf forests to the pluvial
glacial periods, and the reduction of mesophilous flora, with concomitant
development of semixerophilous and thermophilous vegetation, to the arid
interpluvials. These conclusions are based either on studies of
discontinuities in plant distribution, or on the geological literature
(Vereshchagin, 1949c). An interplay of many factors, other than climatic,
is responsible for discontinuities in the distribution of species. Even the
penetration of northern plants (e.g., sphagnum and sundew) far into the
south — to the Greater Caucasus, the marshlands of Colchis and the Lesser
Caucasus highlands — cannot be correctly evaluated unless all the factors
which control species distribution are known. The use of phytocenotic
transformations to detect climatic changes is open to question even when
applied to a period of time as short as the historical epoch (е.5., Sinskaya,
1933; Fedorov, 1952). The Pleistocene climatic changes cannot be evaluated
from the fossil plant record alone because the number of known macroscopic
plants is small. However, the available paleobotanical and geobotanical
evidence indicates that the flora of the Caucasus in the Middle Pleistocene
resembled the Recent flora and the plant habitats had the essential aspects
of the Recent. The arboreal vegetation at the end of the glacial epoch was
also similar in type and in distribution areas to that of Recent time
(Grossgeim, 1936, 1948).
The valley glaciers of the past, as, for instance, those on the slopes
of western Caucasia, were much larger than those of the Recent. This is
81
(8 TABLE 6, Stratigraphic summary of post-Tertiary history of the Black Sea, the Caucasus and the Caspian
Sea (from Vardanyants, 1948, p, 23) and connections via the Bosphorus and the Manych region
Bosphorus Black Sea Manych Caspian Sea
Buhl stages Ореп (?) |Post-Khvalynsk layer
Closed Regression
Subsidence, regression, Uplift, glaciation (Buhl
Closed ;
м decreasing salinity or Neo-Wirm )
Ancient Black Sea beds Open Khvalynsk layer
о Subsidence, influx of Uplift and glaciation Closed Regression
en
ы seawater (Wiirm )
Neo-Euxine beds Riss-Wiirm (?) Khazar layer
Closed Subsidence, regression, Uplift and glaciation Closed Regression
decreasing salinity (Riss )
Karangat beds Open Upper Baku (transitional
beds )
Subsidence, influx of Uplift Regression
Open
seawater
Uzunlarskoe and Ancient Middle Baku
i Closed
Euxine beds Lower Baku
Chauda beds Uplift and glaciation Closed Regression
(Mindel)
ИИ Regression and decreasing] Uplift and glaciation Closed Regression
Os
salinity (?)
indicated by the occurrence of glacial troughs and terminal morainic ridges
in the valleys of the Laba, Zelenchuk, Kuban, Baksan and other rivers in
the area (Figure 41).
The glacial troughs occur in the upper parts of the valleys, far short
of the Skalistyi ridge. It is clear that there were no basic differences in
the environments of the faunal and floral complexes between the time when
the valleys were covered by glaciers, andthe Recent. The vegetation of
the highlands descended, although it was not completely displaced, to the
87 piedmont plains. * During the period of maximum glacial advance, there
was adequate living space for mountain animals on the meridional and
lateral mountain ridges near the glaciers. Even at present, mountain forests
* Even Grossgeim (1936, p, 203), allied as he was with Kovalevskii, was compelled to recognize this fact,
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88
(coniferous, broadleaf and mixed) and mountain animals exist close to the
glaciers, as on the Cherek, Urukh and Rion rivers (Figure 42).
The gravel beds and large boulders in the piedmont plains were more
likely the result of mudflows, caused by floods and volcanic activity, than
relicts of hypothetical glaciations.
The evolution of mammalian faunal complexes must be related to the
snow factor andto the latitudinal shift of the climatic zone on the Russian
Plain caused by the northern European glaciation. However, the depth and
duration of the Pleistocene snow cover have not been studied, and can only
be inferred from examples of distribution and migration of some mammals
in the Recent.
FIGURE 41, Glacial trough and terminal moraine, overgrown with pine, in the Tseya ravine in
North Ossetia
Photograph by author, 1947
The occurrence of boreal plant species in the Caucasus can only be
accounted for by the proximity of the Riss glacier tongues to the Caucasian
Range (Medvedev, 1915; Grossgeim, 1936, 1948). In other words, it is
assumed that tundra and taiga existed in the Manych area and in the Salsk
steppes. Results of recent palynological and paleontological studies indicate
that steppes always existed between the lower Don and the lower Volga
rivers.
We will evaluate the paleontological and zoogeographical data in the
Caucasian stratigraphy after we review the Pleistocene localities. It should
be noted that the hypothesis of extensive continental glaciation in the
Caucasus implies that the evolution of all the highly specialized forms
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89
(mountain goat, snow mouse and other forms) took place within а very short
time — the postglacial period.
The glacial stratigraphic scheme, particularly the hypothesis of
continental glaciations inthe Caucasus, should be reviewed with reference
to biological data. ;
FIGURE 42, Glacier and forest in the upper reaches of the Rion
Photograph by author, 1957
Absolute geochronology of the Quaternary. Application
of the absolute time scale of the Quaternary to the Caucasus did not produce
new results. Most Russian authors accepted the estimates of Soergel (1925)
and Milankovich (1939) of 600,000-650,000 years as the duration of the
glacial epoch. In estimating the time of origin of the Aral-Caspian sea
at 23,000 B.C., of the Khazar sea at 13,500 B.C. and of the Khvalynsk sea
at 3,400 B.C., Kovalevskii (1933) condenses the entire history of man and
the evolution of the Caucasian fauna into a very short period of time. Later
investigators have found that the Quaternary orogenic processes and
glaciations in the Caucasus lasted much longer — 500,000-600,000 years
according to Vardanyants (1948) or even 1,000,000 years according to
Nikolaev (1949) and to Ronov (1949). These estimates, which are of great
importance in the calculation of the rates of evolution, speciation and
development of the fauna, will undoubtedly be amended in the course of
future studies of the physicochemical composition of rocks, bones and
sediments. The only Caucasian stratigraphic stage with a satisfactory
paleontological record is the Upper Pleistocene — presumably the time
when glaciation reached its maximum extent.
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90
MODE OF OCCURRENCE OF FOSSIL MAMMALS АТ
THE PLEISTOCENE) LOCALITIBS
Pleistocene localities are more amenable than those of the Tertiary to
studies of the conditions of animal death and deposition.
In the Caucasus Quaternary mammal fossils occur in marine terraces
along the coasts of the Black, Azov and Caspian seas, in river and lake
terraces, in the surface loams and in caves.
Only a few occurrences of mammals in marine and lagoonal sediments
are known in post-Apsheron strata. This is probably due to the latest
tectonic processes, since it is unlikely that the variability of the animal
world in the Pleistocene was any less than in the Tertiary.
The bodies of animals which had died from various causes (see
Introduction) accumulated at river mouths and estuaries. Smaller
accumulations formed in the alluvial sediments of the middle and lower
parts of rivers. All such sites in the lower parts of the valleys of the
Kuban, Terek, Rion and Kura rivers are at present in the zone of tectonic
subsidence, in which sediments continue to accumulate. Consequently,
most of the sites remain concealed by the sedimentary cover and can be
discovered only in the processes of drilling and quarrying.
Exceptions to this usual mode of occurrence are found in those sites
which were at the head of coastal bays, provided the bays were in the area
of the later uplift of the foothills. Such localities are found in the middle
parts of the Kuban (Kropotkin, Nevinnomyssk), on the Terek River (near
Mozdok and Kotlyarevskaya), on the Rion (at the longitude of Kutaisi and
Zestafoni), and on the Kura (in the Mtskheti-Tbilisi area).
It is possible that several fossiliferous layers occur at localities ina
"Zone of undulation" (i.e., longitudinal depressions and swells) because
favorable hydrological conditions prevailed over several epochs.
Most of the Pleistocene mammals are found as intact or fragmented
skeletons in continental deposits of various origins (alluvial, lacustrine,
diluvial, eolian) in caves, travertines and asphalt pools, on the plains and
in the foothills of the Caucasus Isthmus.
Although life was precarious in the mountains and avalanches, falling
rocks, slippery, ice-covered surfaces and hunger brought frequent death
to the animal communities, the chances were poor for the preservation of
their remains. The fine-grained alluvial sediments of mountain rivers
did not offer much likelihood of preservationeither. Such sediments are
usually thin and tend to shift with time (Shantser, 1951). Fossil vertebrates
do not occur, as arule, in the terminal and lateral moraines of mountain
valleys, because of the attrition of bones in the process of moraine
deposition.
The diluvial sediments at the foot of the mountains, far from the valleys
of large rivers, do not contain fossil accumulations, since skeletons were
usually destroyed during the slow deposition of the sediment.
Skeletons of animals carried by torrential streams onto the plain areas
stood a better chance of preservation in the thick mud and conglomerates
formed by the streams. Mudflows continued to cause destruction and death
among animals throughout the Quaternary; they have been vividly described
by Ivanovskii (1911) for the Araks River valley, and by Muratov and
Gvozdetskii (1948) for the Khasaut valley in the northern Caucasus.
Mudflows are also known to have occurred in the rivers of eastern
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91
Transcaucasia, for example, in the Demir—Aparan-Chai ravine.
Mudflows are produced by such factors as heavy rains and volcanic
eruptions which melted the firn and ice. The flows, which were more
frequent and powerful in the Pleistocene, killed masses of animals.
Heavy snow and surface ice were probably common causes of violent
death among the Pleistocene animals of the plains, just as they are among
the Recent animals. On the plains beyond the river valleys bones were
rarely preserved and then only inthe gulleys and diluvial loams.
Disposal of food remains by primitive man at campsites and in caves
added a new factor to the accumulation of Pleistocene fossils. The prey
carried by predatory animals and owls also contributed to the bone
accumulations of many species at the same cave sites.
This type of death assemblage occurs most frequently in the mountains
and is the main source of information on the history of the mountain fauna.
The sections which follow are devoted to the discussion of the major
localities by regions.
PLES © OG ENE ee ON fb ввАВУМО ОСА rei aes
Western Ciscaucasia
Following the deposition of the bone-bearing Psekups gravels and
conglomerates onthe Taman Peninsula, norich, diversified mammalian ac-
cumulations are known for quite a longtime. The Lower and Middle Pleistocene
fossil record consists mostly of teethof Elephas meridionalis and
№. pPeEooonnt hie ria.
On the Kuban Plain, i.e., on the right bank of the latitudinal section of
the Kuban, fossil mammals occur at great depth in the sediments of ancient
rivers which ran off the Caucasian mountains.
The remains were deposited on the edges of floodplains of the plavni or
liman type fed by mountain streams, probably beyond the zone of deposition
of coarse clastics.
The clays and sands of the Kuban Plain (e.g., those in the Krasnodar
area) contain bones of large Middle Quaternary mammals of Khazar time,
along with wood and freshwater mollusks (Unio, Paludina)
(Pravoslavlev, 1932). The overlying beds are loess loams with
occasional intercalations of volcanic ash. In addition to mammals and
plants, these beds contain terrestrial mollusks (Buliminus tridens
Mill. ), which are indicative of a steppe environment.
Bones of "Воз, Equus, Cervus, Elephas'" have been recorded
by Pravoslavlev and Ansheles (1930) from the 50-m-thick loams near the
Cossack village of Temizhbekskaya.
A number of fossils from other localities were identified by the author
in 1952 in the Krasnodar Museum. These include the teeth and skull
fragments of anearly mammoth, Elephas trogontherii —
E. primigenius, from the village of Dinskaya, and remains of a long-
horned bison, Bison priscus longicornis, from the village
of Krylovskaya on the middle Chelbas. The skeleton was buried in the
silty sand of a stagnant-water basin. The bones were only slightly
permineralized and resembled alabaster in their appearance. Bones of
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93
Bison priscus longicornis and mammoth were preserved under
similar conditions near the village of Skobelevskaya. The bones were
found in a silty marsh bed 16.5mbeneaththe loess. Elongated grooves, the
tracks of boring invertebrates (i riehiomine rawlarvae wal ), occur on the
surfaces of the bones. The occurrences of B. priscus longicornis
indicate the Middle Pleistocene, Khazar age of these localities.
Five nearly complete skeletons of small cave bear (Grigor, 1926),
described by Borisyak (1930, 1931) as Spelaearctus rossicus, were
found near Krasnodar at a depth of 14 m in the sandy loams of the second
alluvial terrace of the Kuban. The death and deposition of the animals have
been attributed to mudflow by Borisyak.
Girei Quarry. The stratigraphy of the alluvial sediments of the
Kuban Plain has been worked out by Gromov (1948) from the sections in
the quarries near the village ‘of Girei and the Kavkazskaya railroad station
in the town of Kropotkin.
Sandy loams and sands 2m thick, and greenish loam 1.5 т thick
with vertical plant roots have been exposed by quarrying in the 10—15-m-
thick Wurm terrace of the Kuban. Crossbedded gravels with sand lenses
and gravel beds up to 3.5 m thick were exposed in the lower part of the
quarry. Bones of mammoth occur at the base of the 3.5-m-thick sequence,
which is underlain by a clay bed 0.8 m thick with plant remains (Figure 43),
and a sand bed 3 m thick. The sand is underlain by gravels containing
bones of bison, deer, and elephant.
The following species occur in the gravel beds (Gromov, 1948):
Beh ace, Usha ар. г оо hve iain Aimar Monee ва ove mats!
HMhwnocenos meme, В. ant lqumiavise Вой pevusicusmer,
Поет cone nts je. wom ise use chem iia tise:
According to the observations of the geologist Kolbutov, the bones of
E. wusti and R. mercki came from the lower gravel bed, which
overlies the clays with lignitized wood. The age of these fossils is close
to those of the Т1газро!' gravel. The mammoths from the sands and gravels
higher in the section are of Riss age, and thus contemporaneous with the
"fauna'' at the Il'skaya paleolithic locality (Gromov, 1948). The material
represents three faunal complexes: 1) post-Khazar with Elephas
primigenius, 2) Khazar with E. trogontherii, 3) pre-Khazar with
E. wusti.
According to Gromov, the age of most of the bones from the Girei quarry
is Khazar, i.e., Early Riss or Riss-Mindel.
Our studies in 1952 of the locality at Girei revealed the following
stratigraphic sequence from top to bottom: surface loams; ancient alluvium,
12-13 m thick; Sarmatian blue clay, 5-6 m of which were exposed by the
Recent Kuban River erosion (Figure 44).
Fragments of large tree trunks, often coated with ferruginous silt,
saturated with water and charred in appearance, are common in the lower
part of the sands and clays at а depth of 10 т. The following trees, which
are characteristic of the present foothills, have been identified: oak
(Quercus sp.), beech (Fagus sp.), and poplar (Populus sp.).
Bones occur in the gravels overlying the wood-bearing sands. Most of
the bones are well-rounded, heavily permineralized and iron-stained.
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FIGURE 43. Gravels and sands (stripped of loam surface) in the Сие! quarry
Photograph by author, 1952
We identified elephant teeth from the Girei quarry, now in the
Krasnodar Museum, as follows: Elephas meridionalis —two molars
with enamel 3.5-4 mm thick; E. trogontherii —three molars with
enamel 2.4-2.8 mm thick; E. trogontherii primigenius — one tooth
with enamel 1.8-2.2 mm thick. The bone material in the teeth of the
meridionalis and trogontherii species is pink-tinged beige and
resembles marl. The state of preservation is comparable in both species.
A tooth of an early mammoth with 8.5 enamel lobes over 10 cm on М? is in
a better state of preservation: in fracture the bone material is white, black-
veined and slightly iron-stained. The collections also include bone fragments
of deer (Cervus elaphus) and a marl-like fragment of a horn stem of
Bison, ci. schoetensackKis
The sandy-gravelly sequence at Kavkazskaya contains an assemblage
consisting of animal remains which have been eroded and redeposited by
lake sediments of different ages. As a whole, the Elephas meridiona-
lis —E. trongotherii complex is possibly somewhat younger than the
Taman faunal complex (see Chapter I). In all probability, there were no
antelope in the younger complex: the bison were larger than the Taman
species, and the deer were less diversified than those inthe earlier complex.
The bones of mammals which are infrequently found inthe upper parts of the
loess and in the fossiliferous soils underlying the loess on the water divides
of northwestern Ciscaucasia are attributed to a later time. The Middle
Pliocene localities in the lake sediments near Krylovskaya and Skobelevskaya
are exceptions to this dating. Mirchink (1936b) has correlated the upper
* The material was lost during the war.
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94
|
FIGURE 44, Section through bone-
bearing gravels and sands in the Сие!
quarry
a — surface loams; b — sandy loam with
gravelly intercalations; с — river sands
with layers of gravels and clays; d —
gravels and sands with mammalian bones
and tree trunks; e — Sarmatian blue
clays. On the right — thickness in meters
part of the loess in Ciscaucasia with the
Wtirm glaciation, and the fossil soils with
the Riss-Wurm interglacial.
The Basal Quaternary of the Trans-Kuban
Plain is represented by the middle part of
the Psekups gravels with Е. meridiona-
lis, Equus stenonis and unidentified
Bos (Gromov, 1948).
The meridionalis and trogonthe-
rii elephant species are usually dated as
the Lower or Middle Pleistocene. These
occur, as arule, in the highest terraces
of the left tributaries of the Kuban. A tooth
from the mandible of Elephas trogon-
therii was found near the village of
Pregradnaya on the Bolshoi Zelenchuk (ZIN
collection).
The Trans-Kuban Plain collections in
the Krasnodar Museum include elephant
molars of a species intermediate between
Elephas trogontherii and
Egor 1 nalenanains. mney ооо емаюже!
is 1.8-2.4 mm thick; it forms 7-8 lobes over
10 cm of the upper surface. The iron stain
on the bones indicates that the remains of
the elephants were buried in fine-grained
river and lake sands, which subsequently
became marshland. As for their state of
preservation, these teeth always look younger
than the teeth of E. meridionalis from
the Psekups and Taman; the dentine and
cement are dense and light beige and
yellowish in color. In the gravel quarry
near the village of Nekrasovskaya on the
Laba, the following forms have been
identified: tooth of an elephant of the type
intermediate between E. meridionalis
and Е. trogontherii; lower molar of
Equus aff. sussenbornensis;
fragment of a horn of the deer, Euclado-
сегиз sp.; anda fragment of a horn of
a giant deer, Megaceros euryceros.
The faunal composition, the mode of
preservation and the iron stain on the bones
at this locality resemble some deposits on
the Taman Peninsula.
More recent collections by the Krasnodar Museum from the gravel
quarry near the village of Ivanovskaya include molars of true mammoths
with enamel 1.4-1.8 mm thick and 12 lobes over 10 cm. A molar ofa
mammoth was also found on the Pshish River. Teeth of Elephas
meridionalis and E. trogontherii from the vicinity of the village
_of Novo-Labinskaya оп the Laba River are in the Maikop Museum collection.
89
95
The museum material is difficult to place stratigraphically because of
inadequate labelling and the peculiarities of the mode of deposition on the
piedmont plain (see above). On the plain, cones of silt debris, formed out
of mudflow and carrying the bodies of animals, were deposited one on top
of the other. However, these deposits were later eroded by rivers which
redistributed the bones in aggradation terraces. In this way, faunas of
quite different ages occur in the same river terraces, and, for this reason,
Mirchink's (1936b) attempted synthesis of the geomorphological and paleon-
tological data on the river terraces of the southern Russian Plain and of
Ciscaucasia can be accepted only with great reservation. On the basis of
the occurrence of Elephas planifrons teeth, Mirchink dated the
175 m terrace near Cherkessk Pliocene in age. The 100-125 m terrace
belongs to the Mindel glaciation period, according to Reingard. On the
Kuban Plain, this terrace is overlain by two loess strata. Two
layers of osseous soil, correlated with the "Ра1а41па sands" near
Taganrog, Bessergenovka and Yeisk, occur here. The 50m
terrace near Cherkessk has been correlated by Mirchink, without
paleontological evidence, with the moraines near the village of Khumarinskoe
and the lower loess horizon occurring on the water divides of western
Ciscaucasia. The loess, according to Mirchink, is of Riss age. It is likely
that the loess in Ciscaucasia correlates with the gravels at the base of the
second terrace in the lower Volga. Fossils of the Khazar faunal complex
occur in these gravels.
Il'skaya Paleolithic site. The activities of Paleolithic man in
western Ciscaucasia are traceable through flint tools which occur on the
shoals of the Kurdzhips and Belaya rivers near Maikop and Fortep'yanovka.
The area probably had many summer encampments. The caves in the
foothills, mentioned in local accounts, have not as yet been investigated
by paleontologists. The Fanagoriiskaya cave, a narrow, cold crevice cut
through the limestone conglomerates by the upper Ayuk rivulet west of the
village of Goryachi Klyuch, was unsuitable for large animals.
The Il'skaya site was discovered in 1898 by Debaille (see Volkov, 1899).
This is an open site on the right bank of the П', the left tributary of the
Kuban, 40 km southwest of Krasnodar. The site, now in deforested foothills,
was rediscovered in 1925 by Zamyatnin (1934) who carried out excavations
in 1927-1928, and who was followed by Gorodtsov (1940) in 1930 and 1936-
1937.
Paleolithic remains occur in the lower of two beds of fossil soils in
diluvial loams located on the second 10-14 m terrace of the П' under the
dolomite cliffs.
The archaeologists mentioned above have described numerous tools found
at the site made of dolomite and, occasionally, of flint. Because of the
poor finish of the dolomite tools, Zamyatnin dated the site Mousterian,
but the more refined finish of the flint tools led Gorodtsov to date the site
Solutrean. The geological dating of the terraces is Riss-Wturm, Early Riss
(Gromov, 1933, 1948).
According to Gorodtsov, the community abandoned the site when the
surface of the Rissterrace became bogged at the onset of diluvial processes.
At the same time oil seepage from the dolomite displaced the water at
the site and preserved the strata containing Paleolithic tools and bones.
The state of preservation of the bones varies at the Il'skaya site. Some
90
36
fragments ''jingle'' and are so thoroughly permineralized that they only
stick slightly to wet fingers; their specific gravity is almost equal tothat of
Pliocene bones. These bones are usually strongly stained with the iron
oxide which fills the pores, or are covered with particles of silty fine-grained
sand of a type characteristic of swampy springs on river floodplains. The
enamel of the teeth is completely black. Those bones which were preserved
in oil are younger in appearance. АП the collected bones are food remains —
the epiphyses on the bones are broken off, and the diaphyses split as though
bone marrow has been extracted. Fragments of jaws and individual teeth
of bison are particularly abundant and bones of young mammoths, showing
teeth impressions of large carnivores, also occur at the site.
When the asphalt which adhered to the teeth of the bison was dissolved
with benzine, the remains of birch mice, small snakes, beetles and grasses
became visible.
Table 7 and Figure 45 give the species and their number in the Zamyatnin
and Gorodtsov collections, as identified by Gromova (1932a, 1937) and by
the author.
TABLE 7. Mammals from the II'skaya site
Number of
Species and higher taxa
р 8 individuals
Number of bones
WS WS, PSPS RATERS celle о, о eA
Canis patiee Шири (simallliform)ins 2) erenalewe
er UIE! DLS а о | rachel TTS) had ls
Cirojeuiatisprenl aera. so асе
ENEMAS Реми © ММЗ оробосбвооосанов с
LS DUR Ai, SMROPACWI a slob с. obc ol
вета vere awicaisicial И. нии
Muridae, not determined below generic level ....
Ее ми) Bawa US) фл Мое. Bile een
Een. я ое che. cane
SHUUS CEO! ати о. oie А ое mR
Gervirsmeblea'p Hiuish- te со оо о о
Щеста сео тему ео; А: се
ЗОО SC WSN Alc) ot AN о Мо с ВИ
S QS el) vara MOE о р Оооо во басовые
Artiodactyla, not determined below generic level
i oS — co cn p> № на = Sh 5
The absence of woolly rhinoceros, elk and mountain goat is probably
not accidental. These animals were rare and difficult to hunt.
Elasmotherium and camel, common inthe Middle Pleistocene of
the Volga River region, either did not exist on the Trans-Kuban Plain, or,
at most, were extremely rare.
Bison remains account for 87% of the bones at the Il'skaya site. The
Species, according to Gromova, is one of the forms showing reduction in
Size ва ет ев‘ deminutus.
91
In fact, relatively small molars and carpal bones (of cows?) occur along
with bones of large bison, which are indistinguishable from those of Khazar
age of the middle and lower Volga. The proportion of large bones is small,
(98)
= Sen
ре
7 eA NY a Е
in
FIGURE 45, Mammal remains from the Il'skaya site
1 — jaw of Canis lupus; 2— jaw of Crocuta spelaea; 3,4—My,of Elephas primigenius
showing worn surfaces; 5 —upper jaw of Sicista cf, caucasica (x8); 6 —phalange of Equus
hidruntinus; 7—metacarpus of E, caballus;
1704 92
97
but the determinant here may have been the selectivity of the ancient Trans-
Kuban hunters rather than any morphogenetic factor. It is possible that
hunters might have killed cows in preference to bison.
The surroundings of the site were probably meadow- and forest-steppes,
which were inhabited by large herds of bison, horse and deer.
The climate and landscape of the Trans-Kuban region in the Upper
Pleistocene can be best inferred from the presence of saiga and boar. Saiga
did not live on the left bank of the Kuban in historical time, because of the
damp summers and snowy winters which prevailed there. Boar, however,
survived until recently by descending to the forested plain after heavy
snowfalls in the mountains. Thus it can be seen from the evidence at the
Paleolithic site that the Trans-Kuban area was more of a steppe than it is at
present; the climate was drier, and there were no catastrophic snowfalls.
However, the saigas could migrate from the north into the foothills only
over ice- and snow-covered ground, as they do now in eastern Ciscaucasia.
FIGURE 45 (continued )
8—jaw of Megaceros euryceros; 9—metacarpus of Cervus elaphus; 10 —horn stem of
Saiga tatarica; 11, 12 —Mgand epiphysis of shank bone of Bison priscus.
93
The negligible number of boar bones is probably accounted for by the
much higher altitude of the mountain forests at that time. The absence of
rock and mountain animals — mountain goats, sheep and boreal mammals —
is also highly indicative.
The absence of mountain goats, however, cannot be used in dating this
site and estimating the extent of glaciation in the Caucasus. Even during
the maximum extent of glaciation, these animals could have lived along the
margins of the mountain glaciers near the Semashkho and Oplenen peaks,
60-70 km from the site, and well outside the regular hunting grounds of the
Il'skaya hunters. The mountains close to the site were low and gently sloped.
The absence of the arctic fox, reindeer and ox indicates, most likely, that
these species did not reach the northern Caucasus.
The elephant bones at the П'зкауа site belong to a mammoth of a late
type (tooth enamel thickness: 1.3-1.4 mm). At least two adults and three
young, at the stage of tooth growth — Me-M‘4, were found.
New collections of the bones of smaller species, which are preserved
at the site in asphalt crusts, would contribute much to the understanding
of the evolution of the Quaternary fauna in the Caucasus.
Fragments of insect chitin which fell into the оП pools at the site and
were thus preserved, were examined at our request by Bogachev, who
identified eighteen beetles, one ant and one wasp (?). Of the beetles, the
most common are water scavengers and water beetles (Dytiscus sp.
noy.g Agabus*spifrboiwsesp., Hydroporus sp., Colym Баев
fuscus L.); others are ground beetles (Carabws sp. nov.ja@t Tat ysma
sp., Amara sp., Bembidium sp.), leaf beetles (Gastroidea sp.,
Donacia sp.) and Coprinae (Aphodius subterraneus L.). The
steppe forms are represented by Tentyria nomas Pall. Many of the
species differ considerably from the Recent forms, which indicates the
antiquity of this fauna.
Among the 20 grasses preserved in chunks of bitumen, collected by
Zamyatnin and studied by Kipiani, the following predominated: Cruciferae
gen. et sp., Compositae gen, et spr, Le pidium) perfoliatum,
Polygonum aviculare mh и р Ног baa ici. pal@etiris, cilagtatice,
Chenopodium sp. etcf. Atriplex. There were also occasional remains
of Aligma sp., cf. Carex) Батрив. sp., Plantaco spijeandsetrer
forms. No macroscopic remains of arboreal species were recorded. The
species listed are ancient, weedlike types, characteristic of sunny habitats
and some of them of a shallow-water environment. Cherdyntsev has set
the age of the Il'skaya site from the carbon-dating of the bones at 39,000
years.
In our opinion, the Il'skaya fauna indicates the beginning of the Upper
Pleistocene and the extensive development of steppes in the foothills region.
In 1957 Formozov carried out exploratory excavations at the entrance
to a cave in the Belaya River ravine near the village of Dakhovskaya. He
collected eighty fragments of poorly preserved bones and some tools of
the Mousterian type from the 3-m level. We have identified the bones as
follows: Canis sp. (10/2), Vulpes sp. (17/4), Ursus spelaeus (3/2),
Crocuta spelaea (1/1), Meles meles (7/2), Panthera spelaea
(4/1),.Felis%ef, Silvestris (2/1), Lepus europaeus (3/1),
Cricétus cricetus (3/2), Elephas cf. primigenius (14/7),
Cervus elaphus (5/2), Megaceros euryceros (5/2), Capra cf.
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100
caucasica (2/1), Bison priscus (19/3). * This fauna is characteristic
of ecological assemblages of the Middle-to-Upper Pleistocene transition.
As a whole, it is quite similar to the Il'skaya assemblage. Some small
forms of wolf and fox are particularly interesting.
Cempenll Cis Came ssi 2
On the Stavropol Plateau and in the Pyatigor'e area Pleistocene mammals
occur mainly in diluvial loams covering the water divides and filling old
gulleys. Older, Lower Pleistocene fossils occur in deposits of ancient
streams and marine and lake bays on the surrounding plains. The Lower
Pleistocene material is represented by Elasmotherium sibiricum
from the vicinity of Divnoye in the Manych depression(Pavlova, 1916), and
by Equus cf. stenonis teeth from loess loams near the village of
Mirnyi (Khomenko, 1913a). The following finds can be dated Middle
Pleistocene, Khazar: hornof Megaceros hibernicus, found by
Khomenko in diluvial clays in the Kalaus cliffs; Saiga prisca
(= S. tatarica) from diluvial clays in the Koronkoyar gulley, 35 km south-
southwest of Stavropol; Equus caballus fossilis, in diluvial clays
of the Shatyr mound, and remains of Bos sp. from sands at the bottom
of a well in the stream bed of the Mutnyanka.
The collection of elephant teeth in the Stavropol Museum includes three
lower molars (7 lobes over 10 cm; enamel thickness 2.5-2.8 mm) of Lower
Pleistocene Elephas trogontherii. The museum collections also
contain six fragments of long-horned bison skull and one fragment of skull
of aprimitive bull (Bos primigenius) which was found in the Stavropol
area. Most of the material was collected from beds underlying the loess
loams and can be dated Middle Pleistocene.
New materials were collected by Gnilovskii in 1941 on the Kuban-
Yegorlyk canal on the western edge of the Stavropol Plateau. The following
specimens were collected from a 6-m depth of the first terrace of the
Yegorlyk: two fragments of mandible, a lower carpal epiphysis and a heel
bone (bitten through by some carnivore) of Paracamelus cf. gigas;
fragmentary bones of short-horned bison; and fragments of neck vertebrae,
horn, forelimb bones and astragalus of Bison cf. schoetensacki.
The bones are chalky in appearance, with light iron oxide stains along the
cracks.
The following bones were collected from a 9-m depth in the alluvial bed
overlying the Maikop clays in the first Yegorlyk terrace: fragments of upper
molars of bulls (Bos sp. or Bison sp.) and an upper molar and first
phalanx of a small horse (Equus sp. ). The bones, which are iron-stained
and not rounded, occur on the erosional surface and are probably Lower
or Middle Pleistocene.
Ryabinin's (1918) collections from the Udel'naya steppe of the upper
Kalaus are of Upper Pleistocene and partly Holocene age. The material
was collected from the upper part of the loess loams exposed in the left
bank of the Malyi Kiyankul' gulley, at the upper end of the Pervaya
Ternovaya gulley, and in the Kalaus stream bed near the opening of the
* The numerator indicates the number of fragments; the denominator, a possible number of individuals,
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101
Malyi Yankul' gulley. At the Malyi Yankul' locality numerous mammal bones
occur at a depth of 2.5 m inthe structureless brown, sandy, gypsiferous clay
with small, limey concretions. Ryabinin has identified the following forms:
"Equus caballus fossilis; ба во scrofarfossilis) Dos taurus
fossilis, Saiga tatarica fossilis anda horn similar to those of
goitered gazelle (Gazella subgutturosa).'' Mesolithic tools (of the
Solutrean or Magdalenian cultures, '' according to Ryabinin) collected at
the locality indicate the last part of the Pleistocene or even the transition
to the Holocene.
Thus, the fauna of the margins of the Stavropol Plateau does not support
the hypothesis that mountain mammals of the Caucasus migrated to the
plains during the maximum glacial advance. No boreal species have been
recorded in the area. Studies of the caves on the southern and western
slopes of Mount Strizhament (Vereshchagin and Gromoy, 1953b) would
contribute to the knowledge of the steppe and forest-steppe complexes of
the Stavropol area of the Upper Pleistocene.
Bones of a mammoth have been recorded from gravel quarries near
Yessentuki in the region of the Pyatigor'e laccolith on the Kabarda sloping
plain (Karpinskii and Dreier, 1904). Eikhwal'd(1850, p.190) reported
finding near Kislovodsk the complete jaws of a primitive form of Equus
caballus covered with tuff.
Travertines of the Pyatigor'e area. Travertines were
deposited in post-Pliocene Pyatigor'e by mountain springs, which changed
course after each mountain-building cycle. Animal remains were shifted
and scattered by carnivores over the slopes until they became covered by
sediment. Layers of lime-tuff formed on the bones upon contact with
water rich in calcium-carbonate.
Travertines were deposited on almost all the laccoliths — Lysaya,
Zheleznaya, Zmeika and Beshtau. However, only the travertines of Mount
Mashuk have beer subjected to paleontological study (Ivanova, 1948;
Mirchink, 1936b; Gromov, 1948).
According to Ivanova, there are four stratigraphic complexes of
travertines: the first of Pliocene age (see Chapter I), the other three of
Pleistocene and Holocene age.
The travertines of the second complex (Mindel and Mindel-Riss age)
occur in patches on the northern and eastern slopes. Teeth and jaw
fragments of Elephas aff. antiquus have been found in these
travertines. The travertines of the third complex cover a larger area to
the northeast. These travertines have been correlated with the 55-m Riss
and Riss-Wurm terrace of the Podkumok, where numerous bones of deer
(Cervidae) occur. The fragments of Lepus sp. and Capreolus sp.
teeth, collected by Kobutov in the Boichenko quarry, probably come from
the travertines of the third complex. The material is Upper Pleistocene
in appearance.
The travertines of the fourth (Upper Pleistocene to Recent) complex occur
in the first and second terraces on the southern slope of a mountain near
Pyatigorsk, which extends to the stream bed of the Podkumok. According
to Gromov (1948), the teeth of an advanced type of Equus sp. and of
Elephas primigenius found inthe gravels and loams of the second
(Wurm) terrace originally came from the travertines.
96
The Pleistocene collections in the Pyatigorsk Museum include a sample
of dolomitized travertine with a calcitic mold of a horn of a small bison
(Bison sp.). Also in the museum is a fragment of the upper jaw ofa
mammoth with the third milk tooth, collected from quarry No.1 on the
northern slope of Mount Mashuk. The fresh appearance of the bone is very
unusual among fossil findings in the south and suggests that the Pyatigor'e
mammoth died as recently as the Holocene. The fragment of mammoth
upper molar found in the travertines near the Medovyi waterfall in the
102 vicinity of Kislovodsk, the mammoth teeth found in the sands and loams
near the village of Kamenskaya, and those found at a 12-m depth in the
banks ofthe Yutsa River are all older in appearance. The museum
collections also include two jaws of woolly rhinoceros (Rhinoceros
tichorhinus) and six skull fragments of primitive bison. The state of
preservation of this material is Upper Pleistocene.
All these fossils from the alluvial and diluvial sands and loams of the
sloping Central Ciscaucasian Plain give only a very schematic picture of
the ecological assemblages of the mammoth fauna which lived near the
mountains. Inthe Lower Pleistocene the region was undoubtedly a steppe. At
the end of the Pleistocene, however, the region became forested, at least
in Pyatigor'e as indicated by occurrences of deer and gazelle bones in
the travertines of the third complex of Mount Mashuk, апару the occurrences
of post-Pliocene ''flora.'' Krasnov has identified 14 species of grasses
and trees from the travertines of the Pyatigorsk area — species which are
also characteristic of Recent broadleaf forests of the foothills: beech
(Fagus sylvatica), filbert (Corylius avellana), white beech
(Carpinus betulus), elm (Ulmus campestris), oak (Quercus
robur), Norway maple (Acer platanoides) and other forms. The
"flora'' from Mount Zheleznaya, which is similar in composition, though
somewhat later inage, was identified by Palibin (1913) as follows: willow
(Salix alba and other species), linden (Tilia cordata), ash
(Fraxinus excelsior) and medlar (Mespilus germanica). No
boreal elements have been recorded at these Upper Pleistocene or Lower
Holocene localities; on the contrary, the southern elements are more
abundant in the floras.
Eastern Ciscaucasia
Most of the plain in eastern Ciscaucasia was covered by the Caspian
transgression, which accounts for the scarcity of Pleistocene mammals
in the sediments of the plain.
The semideserts and steppes in the lower reaches of the Kuma, Terek
and Sulak were in the zone of subsidence. The postglacial marine and
river sediments of the area are at present below the base of river erosion.
As a consequence, the only known fossiliferous localities with Pleistocene
mammals are situated farther west оп the high terraces along the shores
of Pleistocene bays, on the piedmont plain and in the foothills.
The sloping plains of eastern Ciscaucasia are covered by diluvial
sediments and the alluvium of the Terek, Sunzha, Sulak and other rivers.
The relief of this area is more complex than that of western Ciscaucasia
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10
аз а result of uplift of the Terek and Sunzha ranges during the Tertiary and
erosion and further peneplanation during the Pleistocere. These ridges
now separate the Vladikavkaz [Ordzhonikidze] and Sunzha plains from the
alluvial plains of the western Caspian region.
As early as 1886, Pohlig mentioned mammoth bones from the Sunzha
valley in the Caucasian Museum collection in Tbilisi. Mullaev (1929)
described a tooth of Elephas trogontherii from the post-Tertiary
beds on the Ursdon River near the village of Karagach in the southwestern
corner of the Vladikavkaz Plain. Scapula fragments of E. trogontherii
from the 5-m level of the Mozdok sand quarry are in the collections of the
Pyatigorsk Museum. Teethof E. trogontherii of an early type from
the Grozny Region are in the collections of the Grozny Museum.
All the fossils came primarily from Lower Quaternary alluvial sediments.
Gromov (1948) recorded two skulls of Bison priscus deminutus
and a fragment ofa Megaceros sp. skull from the first terrace of the
Terek near Mozdok. The cranial cavities of the bison skulls were filled
with volcanic ash. This fact establishes the time of the fossils as the time
of deposition of the volcanic ash which occurs under the gravels of the
first terrace and on the face slope of the second Riss terrace on the
eastern Slope of Mount Mashuk.
It seems possible that, in the last outburst of volcanic activity of the
Kazbek and Elbrus in the Upper Pleistocene, animals were sometimes
killed and buried in the ash deposited by torrents. Of the fossils from the
Kuma valley below Budennovsk, a lower molar of a mammoth is included
in the collections of the Pyatigorsk Museum. The Upper Pleistocene fossils
are represented by fragments of Sus scrofa ferus mandibles collected
by the geologist Plamenevskii in the alluvial sediments of the first (Wurm)
terrace on the right bank of the Terek above Ordzhonikidze.
Generally speaking, the evolution of landforms and faunal complexes
was similar in eastern and western Ciscaucasia. The same large mammals
occur in the Lower and Middle Pleistocene —Elephas trongotherii,
giant deer, primitive bison and — in the Upper Pleistocene — boar.
Forests were probably well developed in the Upper Pleistocene along
the lower northern slopes of the Caucasus, and even inDagestan. Whitebeam
(Sorbus aria) and European hornbeam (Carpinus betulus) have
been identified by Palibin (1913) as ''post-Tertiary'' species from the
foothills near Makhachkala. These species also occur in the area at the
present time.
Very few fossil mammals are known from the piedmont and inner parts
of Dagestan, or from the marine terraces of the western coast of the
Caspian Sea, from Sulak to the Apsheron Peninsula. The collections in the
museum at Makhachkala contain tooth fragments of Elephas trogon-
therii of an early type (enamel thickness 2.5-3.0 mm) from the
ferruginous gravels of the Manas River. A tooth fragment of a more
recent elephant, with features of Elephas primigenius, and the atlas
of an elephant from the vicinity of the village of Unty are in the same
collections.
Investigations of Paleolithic sites and excavations of caves and sheltered
areas in the Apsheron limestone above Makhachkala and inner Dagestan
would contribute to a better understanding of the evolution of Middle and
Upper Pleistocene fauna in this part of the country.
98
104
Western Ciscaucasia
In western Ciscaucasia the Pleistocene mammals occur mainly in cave
deposits, kitchen middens and food remains of carnivores.
Other types of deposition are seldom found. Several factors account for this:
a lack of sedimentation and arapid stream-flow which destroyed the bones,
a humid climate which induced rapid decay of bone material, and the rapid
subsidence of the sea bottom in coastal areas which precluded the formation
of deltas.
Some occurrences are known from terraces and from small reed and
sphagnum marshes and lakes which still exist in the region of Sochi,
Pitsunda, and Ochamchiri. Some of these localities were later uplifted and
exposed by river erosion. There are extensive alder and sphagnum marshes
along the lower Rion and Supsa (Poti, Kobuleti) in the area of the Colchis
depression which is at present rapidly subsiding under the load of river
deposits. Without artificial excavation, whatever fossil mammals occur in
those beds will remain there for centuries to come, practically inaccessible
to study.
The tooth of Elephas primigenius Blum. in the Sochi Museum
(mentioned by Gromov, 1948) might have been introduced into the area.
Ryabinin (1937) has described a fragment of the right half of the mandible
of a large cat, similarto Felis cf. spelaea, found in 1935 6 km north
of Sochi. The mandible was taken from the gravels of the third marine
terrace (Karangat) and originally dated Riss-Wurm. More recently it
was identified as Early Riss (Vardanyants, 1948). Shells of Cardium,
Tapes and Mactra occur in the same terrace.
The Black Sea coast of the Caucasus was favored by Paleolithic hunters
for its propitious conditions for big-game hunting: numerous narrow valleys
and canyons, coastal terraces which carried the free migration of ungulates
both from the north and from the south, and many caves inhabited by bears.
The uplift of the spurs of the Bolshoi Range in the Pleistocene saved
the cave deposits from river erosion. Paleolithic sites occur along the
foothills from Sochi and Tuapse in the north to Kutaisi and Batumi in the
south and the east. Paleolithic deposits probably continue along the northern
coast of Asia Minor.
Zamyatnin (1937a) has described and mapped 43 Paleolithic localities in
Abkhazia. He subdivided the sites into three chronological groups: 1) pre-
Mousterian, 2) Mousterian, 3) Upper Paleolithic. Most of the sites,
however, are represented by only a few flint tools.
Efimenko and Beregovaya (1941) have listed 15 open sites and 14 cave
sites in western Georgia.
Fossil mammals are known only from cave deposits in the gorges of the
Khosta, Kudepsta, Mzymta, Tsebel'da, Rion, Kvirila and other smaller
rivers.
Caves on the Black Sea coast
In 1936-1938 Zamyatnin (1940) excavated the Navalishinskaya and
Akhshtyrskaya caves in the Sochi and Adler area in the Krasnodar Territory.
oF
105
Navalishinskaya cave is near the village of Navalishino, 12 km
from the sea, on the right bank of the Kudepsta. Two cultural layers were
found inthe cave: the upper layer (Upper Paleolithic) with bones of hamster
and redeposited bones of cave bear; the lower layer (Mousterian,
Aurignacian), with bones of elk, goat, hamster, cave bear, wolf, birds
and shells of Anodonta and Helix. According to Zamyatnin, the cave
was merely a seasonal shelter of early man.
The mammals from the cave were identified by Gromov (1948), as shown
in Table 8.
TABLE 8. Species and number of bones of mammals from the Navalishinskaya cave
Middle Upper
Species nee.
Paleolithic Paleolithi >
Camis ре ME Tas ss
Я реа CNS favs) А ИЕ ФЕИ atte tate rele
Е CLiuGEtusie В оо сбаь пос
ENING С оо ЕО бою нон
ноша
The Shirokopokosskaya cave (which Gromov called Navalishinskaya II)
is located near Navalishinskaya I. It contained Upper Paleolithic tools,
cave bear bones and a few bones of red deer (Gromov, 1948).
Bones of cave and European brown bear were excavated in 1946 in the
Vorontsovskaya cave in the Khosta ravine (collection of the Krasnodar
Museum). The state of preservation of the cave bear bones is not uniform:
in fresh fracture the mandible bone is brownish and free of organic matter.
The radius and femur are dark brown, and show little loss of organic
matter; when scraped, they exude an odor of fresh bones. It seems that
the cave bear lived in postglacial times.
Akhshtyrskaya cave is located on the right bank of the
Mzymta River 100 m above the river bed at the point where the river leaves
its narrow canyon, near the village of Akhshtyr, 15 km from Adler (Figure
46), and is, at present, inhabited only by bats (mostly horsehoe bats). The
entrance to the cave faces south and a corridor 4.35 m high, 2 m wide and
over 100 m long leads to the cave, which was probably much lower in
Paleolithic time. In front of the corridor is an overhead ledge and two
entrances with two platforms. The eastern platform has been excavated
by Zamyatnin (Figure 47). From the west the cave can be approached
over a narrow, easily defended trail on the cliff. Zamyatnin assumed that
it was intermittently inhabited by man over several millennia. Because of
the elevation of the platform dwelling site, the inhabitants probably threw
away most the bones and tools into the river. In the intervals
between man's occupancy, bears and bats lived in the cave. Two narrow
crevices, which were also inhabited by bears, are located farther west
ana down the cliff.
100
106
According to Gromov (1948) the big cave was formed during the formation
of the 60-80 m terrace on the Black Sea coast, which was raised 112 т
near the Akhshtyr anticline.
FIGURE 46. Mzymta gorge below the Akhshtyrskaya cave
Photograph by author, 1954
Two phases can be distinguished in the geological history of the cave.
In the first phase two perpendicular faults formed: one, a normal fault,
the other, a displacement along the bedding planes of the limestone. These
two faults resulted in the formation of the cave which then, in the second
phase, was continually enlarged by dissolution of the ceiling. During this
period detritus, loam and fragments of stalactites continued to accumulate
on the floor.
Gromov assumed that detritus accumulated with cooling of the climate,
whereas decomposition of the detritus into loam occurred in a warmer and
more humid climate, but the faunal data which follow do not support these
assumptions.
Zamyatnin (1940) has identified five cultural layers in the cave from the
Middle Paleolithic (Lower and Upper Mousterian) to the Recent. He
correlated the Lower Mousterian of the Akhshtyrskaya cave with the beds at
the Il'skaya site in Ciscaucasia and with the Kiik-Koba cave in the Crimea.
The excavations produced over 3,000 identifiable fragments of bones,
mostly from the Middle Paleolithic layers. The material includes bats,
carnivores, rodents, ungulates and birds.
Remains of large animals, fragments of epiphyses and diaphyses, are
typical food remains. Bat bones and excreta have accumulated in the cave
101
into present time. Eagle owls probably carried in the hamster bones which
were found in the cave.
Most of the cave bear bones are metapodia, phalanges, isolated teeth and
107 fragments of limb bones. Exploratory excavations in the sheltered section
of the cave produced complete bones: femora, tibiae, fibulae, humeri,
radii, ulnmae, and skull fragments of bears which probably died of natural
causes.
Й
|
1
| Zamyatnin's
|
|
excavation
FIGURE 47, Diagram of the frontal part of the Akhshtyrskaya cave
The matrix on the bones of Mousterian and Upper Paleolithic specimens
is indistinguishable to the eye, but specimens in varying states of
preservation occur within the same bed. The bones are pinkish yellow to
light brown in color. The enamel of the teeth is either the natural color
or darker with black cracks, and the rootsare yellow, brick or brown.
The Mousterian bones are darker, with some surfaces spotted with dendrites;
numerous dendrites may have coalesced resulting in a dark brown color
over the entire bone. In fracture, the bones are usually light beige. They
are light in weight and free of permineralization.
The composition of the mammalian fauna of the cave is given in Table 9
and Figure 48 (author's identification).*
* Our count of bones and individuals in the ZIN collections differs somewhat from Gromovya's: bones from the
Lower and Upper Mousterian were combined, since there is no noticeable difference in the composition,
number and type of preservation,
102
108 A cervical vertebra of an eagle was found in the Mousterian beds. Bones
of the goshawk Astur gentilis L., of the magpie Pica pica К. and
of small sparrows (Passeriformes) were collected from the Upper
Paleolithic beds.
FIGURE 48. Mammal remains from the Mousterian beds of the Akhshtyrskaya cave
1 — metacarpal of Canis lupus; 2—shankof Vulpes vulpes; 3, 4 — canine and astragalus
of Ursus spelaeus; 5 —femur of Martes sp,; 6 —shank of Cricetus cricetus;
7 — carpal of Sus scrofa; 8—second phalange of Cervus elaphus; 9 — first phalange
of Capreolus capreolus; 10 —carpal of Alces alces
The number of bones in the Upper Paleolithic beds is much smaller than
in the Mousterian beds. The smallest possible number of cave bears is 39,
as estimated by the number of phalanges. This was evidently a species
easy to hunt in the canyons where it lived. The European brown bear, is
represented only by a molar from the Upper Paleolithic.
Next in descending order of quantity are mountain goat, deer and boar.
Remains of mountain and rock ungulates are fairly rare: the evidence
of mountain goat has been mentioned, but there is no evidence of chamois.
103
109 The Upper Paleolithic shows a slight increase in the number of boars,
which may indicate not only an increase in species population in the vicinity
of the cave, but also the gradual extinction of the cave bear, which forced
the hunters to look for other game. *
TABLE 9, Mammals from the Akhshtyrskaya cave
Middle Paleolithic
number of
bones
Upper Paleolithic
number of number of
bones
Total
Species individuals
number of
individuals
individuals
Rhinolophus ferrum-equinum
Vulpes wipes aa we о Ose ЕВ
Сап1$ Парики лен
UTsws атс о... ea. <n eee go 37
We ОР ИЕХЯН G6 Seiad .
Мате chy полна о. о
Не СЕ ЗЕ: ое neous ь
Cricetus cricetus
505 зева 4..... о. My
АЛ се$. AGES несе ооо
С ету ще Тарьи д... ес
Megacepos ецгусего № =.
Сартео1 $ сартесо Тиль.
BisOn руси .... АЕ ©
Сарта Galacasica .. Wire. seen
ннюн
©
co
@ tees je ee RS pe)
The absence of wild sheep in the Upper Pleistocene of the Black Sea
coast supports the notion that forests and mesophilous vegetation covered
the area, as inferred from the composition of the Akhshtyrskaya ''fauna."'
It seems unusual that bison and giant deer bones were found only in
Mousterian beds, since the bison has survived in this region to the present.
Elk and gazelle appear only in the Upper Paleolithic beds, though these
species existed in the Caucasus at least from the time of the Lower
Pleistocene. Gromov's (1948, р. 262) theories of climatic changes which
he predicated on the presence or absence of these species are, therefore,
untenable. The number of specimens of ungulates and carnivores (except
for the cave bear) is too small to furnish a basis for sound conclusions on
faunal and climatic changes.
The paleogeographic study of the animals from the Akhshtyrskaya cave
must include the ecology of the species and the hunting traditions of the
primitive tribes.
The highly diversified terrain and the forests of the Black Sea coast
were not Suitable ground for mammoths, horses, giant deer, saigas,
* Gromova (1948) has identified a lower metatarsal epiphysis and the second phalanx of a ram as the small,
wild "Ovis cf, ophion," Further examination shows that the bones are contemporary Holocene in
appearance and their identification as Mousterian is incorrect; they are probably bones of a domestic
sheep which fell into the excavation from post-Paleolithic beds,
104
110
primitive bison and steppe bison. The species occurring near the dwelling
site were hunted for food.
The topography of the Black Sea coast in the Pleistocene was such that
the vertical zonation of the vegetation remained stable, despite climatic
changes. This accounts for the homogeneity of the fauna in the
Akhshtyrskaya cave throughout the Middle and Upper Paleolithic.
The assemblages of mammals from the Navalishinskaya and
Akhshtyrskaya caves are not universal. Nevertheless, they clearly indicate
the existence in the western Caucasus of a boundary between the distribution
areas of the steppe and forest-steppe species of the Russian Plain. Only
the eurytopic hamsters crossed this barrier. No northern species are
recorded from the western Caucasus and the northwestern foothills; nor
were there any southern elements from the dry upland steppes of Asia Minor.
Farther south along the Black Sea coast, Pleistocene mammals occur in
the caves near Sukhumi (Krongel'm, 1873). They were recently discovered
in the Ablaskira cave and in the grottoes in the Tsebel'da canyon.
Ablaskira cave is located on the western slope of one of the spurs
of the Panavskii (Kodor) ridge, 2 km northeast of the village of Otak and
35 km from Ochamchiri. The Achkhitizgo River flows out of the cave
entrance at an elevation of 265 m above sea level.
Paleolithic remains occur in wall niches not far from the entrance. The
cave has been described by Satunin (1910) and Maruashvili (1938b).
Seventy-five fragments of bones of two adult cave bears and one young
were collected in the cave by Sadovskii in 1946. The bones were collected in
the section 50-400 m from the entrance in the clay and gravel of the stream.
The material consisted of tibiae, humeri, metapodia, phalanges anda
nearly complete cranium of a very short-snouted adult specimen. Like the
animals found in the deeper part of the Akhshtyrskaya cave, these died from
natural or accidental causes, rather than at the hand of man. In fracture,
the bones are chalky white; they disintegrate easily under pressure, are not
permineralized and their surface is free of dendrites. There are no data
on the conditions of bone accumulation, the time of deposition of the clays
on the cave floor or the time of human settlement in the cave.
The grottos near the village of Tsebel'dinskoe in the Tsebel'da ravine
were surveyed by the archaeologist Solov'ev in 1941. Paleolithic tools
and bones collected in the Kholodnyi grotto and in Kei-Bogaz are now
housed in the ZIN. The animal fossil material consists of fragments of
teeth and of tubular and plate bones, which were transported partly by
Paleolithic hunters and partly by eagle owls. Identification of the species
in the collection is given in Table 10 and Figure 49.
The state of preservation suggests that most of the bones are of Upper
Pleistocene age; only a small part of the collection (the rodent bones
in particular) may be of Holocene age. The bones represent the local
faunal complex of mountains and forests. The southern limit of the common
hamster is in the area of these southern caves.
Caves of Imeretia
The northeastern part of the Colchis Plain is a low plateau composed
of limestones overlain by Oligocene sands and silts. The tributaries of
105
TABLE 10, Mammals from the Paleolithic sites at the Kholodnyi grotto and Kei-Bogaz
Number
of bones
Number
Species
of bones P
Species
озна, ‘Spe LAVeUS Show, tension Мос SWS, SOTO Ta ye Pa esses ее а 75
Чате RCo в а Cer irsiee lapiitis: eee. ey eee ee 60
маза Vy TA Ва: оли Gapraieeaulcasicae пля o 58
Fielisi sil west mis, пои Pe. wa Вирт С EMM tere Ble 8
FIGURE 49, Mammal remains from the Kei-Bogaz grotto
1 — humerus of Martes зр.; 2— humerus of Felis silvestris; 3 —humerus
of Cricetus cricetus; 4— femur of Arvicola terrestris; 5 — astragalus
of Sus scrofa; 6—second phlanx of Cervus elaphus; 7 -— first phalanx
of Rupicapra rupicapra; 8 —epiphysis of metacarpal of Capra caucasica
the Rion cut canyons 50-60 m deep through the plateau. Over several
millennia numerous caves (dated Upper Paleolithic) were formed in the walls
of these canyons, and were inhabited by prehistoric hunters, cave bears
and eagle owls.
112 The first discoveries of the Paleolithic in western Transcaucasia were
made in the first half of the last century. The Swiss Favre collected stone
tools in 1863 from "Jason's Grotto'' in the Sakazhia cave near Kutaisi
(Zamyatnin, 1957).
106
FIGURE 50, Entrance to the Sakazhia cave
Photograph by author, 1954
Burnt and split bones were collected by the engineer Bernatskii (1884)
in one of the caves near the village of Rgani in the Chiaturi area. Some of
the bones were sent to Rudolf Virchow in Berlin, and some were left in
Tbilisi, wherethey were studied by Anuchin (1887). Bernatskii's collection
consisted mostly of bones of cave bear.
Zamyatnin has dated allthe Imeretian cave sites investigated thus far as
Upper Paleolithic. The earliest sites were uncovered by Krukovskii in the
Chiaturi area: the Khergulis cave near the village of Vachevi and the
overhang, Таго-К1ае, nearthe village of Shukruti. The ''3rd 1ауег" in the
Khergulis cave contained Mousterian and Upper Paleolithic flint tools, and
split bones and teeth of cave bear (Ursus spelaeus), horse (Equus
caballus), Caucasian goat (Capra cf. caucasica) andbison (Bison
ef, prise ws):
Krukovskii's exploratory excavation at the Taro cave produced Upper
Paleolithic cutting tools, Mousterian points, teeth of cave bear, bones of
107
113
Asia Minor hamster (Mesocricetus aff. auratus) and teeth and split
bones of Caucasian goat and bison. *
According to Zamyatnin, the middle group of the Paleolithic sites includes
the following caves: Sakazhia, Uvarovo, Bnele, and Devis-Khvreli.
The Sagvardzhile cave on the Dzevrul rivulet belongs to the same group,
although only preliminary data have so far been published on the many
cultural layers of this cave (Kiladze, 1953; Gzelishvili, 1953).
Sakazhia cave is оп the left wall of the ravine of the Tskhali-Tsiteli
rivulet (the right tributary of the Kvirila), near the Motsameti Monastery,
3 km from Kutaisi. The cave, which was formed near the surface of the
faulted limestone plateau, is now approximately 300 m above sea level
and 50 m above the present stream bed. The arched entrance is 5 m high,
broadening at the top, and leading into a corridor which turns left
(Figures 50, 51).
Even though few of the earlier boxtrees and hornbeams remain, the
setting of the cave against the cascading river and fallen rocks is still very
picturesque. The cave dwellers could hunt their game on the marsh plain
of the Rion and in the nearby mountains.
Indications are that these caves were not inhabited by eagle owls or
bears, except possibly, in the latter case, for very brief periods.
WAI
Ss \Y
= Ms а
х
[>
oe
IID
eel |
\> > IN
SS
YY )
\\
\\\\\ Е Х &
: FZ
SA : Е]
А
AFA РА АХ ЕРАЗ
FIGURE 51, Diagram of the Sakazhia cave
* The bones have been identified by the author from drawings by S,N, Zamyatnin.
108
The Sakazhia deposits are 2 ш thick. The cave was excavated in 1914
by Shmidt and Kozlovskii who dug in the straight corridor, and in 1936 by
G.K. Nioradze. In 1954 the author collected some bones and flint tools at
the far end of the cave where some deposits still remain.
The several thousand flint tools and fragments, which include many
cutting edges and knifelike tools (Figure 52), establish the age of the culture
as Solutrean (Nioradze, 1953; Zamyatnin, 1957). The collection includes
many points and arrowheads made of the diaphyses of horse and bison
114 metapodia. It seems strange that the dating of the thick cultural layer is
FIGURE 52, Sakazhia cave flinttools (from Zamyatnin, 1957) and bone tools (from
Nioradze, 1953)
109
115
so narrow, since the cave was undoubtedly used as а habitation into post-
Solutrean time. Bones of domestic animals which occur in the collection
are probably derived from the upper layers.
Upper Paleolithic food remains are represented by teeth, phalanges,
jaw fragments and epiphyses and diaphyses of skeletal bones. There are
472 specimens in Shmidt's collection, 130 in the author's and over 16,000 in
Nioradze's. *
Most of the bones, however, were not identifiable, because they were
split into fragments. Single bones (heel bone and radius of Felis
spelaea) bear the tooth impressions of large carnivores. The enveloping
matrix suggests Upper Pleistocene age. The fragments are mostly gray
with a slight yellow, free of dendrites and fossilized, but not permineralized
The tooth enamel is a natural color with only occasional darkening. The
canines of a European brown bear appear to be younger — of Holocene age.
Belyaeva (1940c) and Сготоу (1948) published lists of 18 species
identified from a few teeth and phalanges in the Nioradze collection. The
results of the author's studies of both collections are given in Table 11.
TABLE 11. Mammals from Sakazhia cave
Number of
individuals
Number
cgi - of bones
Vulpes vulpes
Canis lupus
Ursus spelaeus „в.
WU. ALC OSD’: @ here Sees GOR © ole ЩЕ. + - вы
Panthera spelaea
Felis ef, Kyinx
Martes cf, foina
Melies meles .. а... See. све
Chionomys roberti
(GAS OW Цени
босс Me 2 © о ме ole
Нузвих ch, Поти ОЖ...
E quis са а И. г
Е. hidruntinus**
pire, Se © слои
Meee ов О Па сс
Sr Orr rP FP rR FP ww Ww PDP
Sus scrofa
Cervus elaphus
a
>
бя Ч о ооо о МАЙ, ее © ое
© © © 0 що ol me. © ца чи №
Capreolus capreolus
А1сез aces, 20m. « «+ о emer о
BiVSiON PiU SIGs, ое see .
Capra caucasicat
Rupicapra rupicapra
* Shmidt's collection is kept in Odessa, Nioradze's in Tbilisi,
** The дата on occurrences of these two species are taken from Gromovy and Belyaeva, since they do not
appear in any available collections.
t In this and the following tables of Imeretian deposits, the goat species is represented by the west Caucasian
form, rather than the east Caucasian, contrary to erroneous identifications by Gromov (1948) and
Gromova (1948),
110
118
119
In addition to the bones listed in the table, ungulates and carnivores
of intermediate size (goat, boar, chamois, wolf) are represented by at
least 8,800 fragments of diaphyses, and larger animals (bison, deer,
horse, cave bear, Panthera spelaea Goldf.) by at least 6,000 fragments.
A few fragments of human bone were also found in the cave.
The diversity of the landscape was reflected in a highly variegated species
composition: bison, goat, deer, horse, boar and cave bear. Because the
cave was advantageously located between mountain and plain, the Sakazhian
dwellers were able to hunt boar, elk and beaver on the Rion marsh plain,
bison (their chiefprey), deer and horse on the Imeretian Plateau, and goat
and chamois inthe rocks of the Rion ravine.
In the Uvarovo and Baratashvili grottos inthe same
canyon, Shmidt collected nearly 30 teeth of Caucasian goat, 12 teethofbison,
12 bone fragments of cave bear, 2deerteeth, 1 horse tooth, 1 fibular tarsal bone
of a wolf, 1 fragment of the pelvis of a beaver and a number of Upper
Paleolithic tools. The state of preservation of this material is similar to
that of materialfound in the Sakazhia cave.
The Bnele cave collectionconsists of afew bone fragments, identified
by Smirnov (1923-1924), together with bones from the younger site at the
Gvardzhilas cave.
The Devis-Khvreli cave is located 4 km from the Kharagouli station
on the right slope of the Chkherimela rivulet ravine in the Cenomanian
crystalline limestones. The cave entrance, which faces southwest, lies
340 m above sea level and 80 m above the Chkherimela rivulet bed. In 1926
Nioradze excavated a section of the cave 25 т long, 4.5m wide and6.5m high,
which revealed four layers: 1) upper layer, with Recent animals; 2) brown
layer, with animalremains; 3) culturallayer; 4) clayey bed. Nioradze (1933)
has identified the site as Upper Aurignacian, whereas Zamyatnin dates it Upper
Magdalenian. The bone collection consists of food remains and bones in
eagle owl pellets. The species, identified by Gromova (1929) and
M.V. Pavlova, are listed in Table 12. (The number of bones available to
Pavlova is unknown. )
TABLE 12, Mammals from the Devis-Khvreli cave
Number Number
РГ of bones SDE cics of bones
аи Ир <5 Ge2 Bula too о: сСартео и: саршео и... .. 2
5 о ре Гаде № ое мани. № АСЕ! AMG ее ес 1
eating а тео Зе, ое ВОЗ ОИ ВО м [ро бони Оо 5 Зо ?
Mesocricetus cf, auratus Cap rz Tei, CAWCASTOE 5. sie ails 22
SPUISHES\C LOR a” Si RMAs ИН. Uj) TOA FA) wa CAINE о. 3
Ce ee
As a whole, the fauna resembles the fauna of the Sakazhia cave. The
smaller number of species recorded is due to the smaller scale of
excavation. It is unusual that no equine bones should have been recorded
at a site located in the center of a dry plateau. Near the spurs of the Rachin
ridge are sites belonging to intermediate and late faunal complexes which
are described below.
111
(116)
FIGURE 53, Mammal remains from the Sakazhia cave
1,2—caninesof Vulpes vulpes; 3,4—canines of Meles meles; 5, 6— jaw of
Martes sp.; 7—Pm3Felis lynx; 8—canine of Felis silvestris; 9—Pmy,and
10 — canine of Panthera spelaea; 11—M, Canis lupus; 12, 13 — metacarpal
and metatarsal of Ursus spelaeus; 14 — jaw (x2) and 15 — first molar (x 10) of
Microtus cf, roberti; 16, 17 — Pmg and its worn surface from Equus caballus;
18 — Мз Cervus elaphus; 19 — Мз Rupicapra rupicapra; 20 — М; Capra
caucasica; 21 — М; Bison priscus;
112
M, ia
un”
FIGURE 53 (continued)
22 — first phalanx of Sus scrofa; 23,24 — first phalanx and astragalus of Cervus
elaphus; 25, 28-30 — horn, astragalus, metacarpal and metatarsal epiphyses of
Bison priscus; 26, 27 — first phalanx and astragalus of Capra caucasica
113
og
if, egy
ye %
yan
See
alt
In 1934 Zamyatnin (19375) collected Upper Paleolithic tools from beneath
limestone overhangs on the right bank of the Kvirila near the village of
Mgvimevi in the Chiaturi area. In 1940 Kiladze (1944) continued excavating
under one of the overhangs.
The bones from Mgvimevi are highly fragmented and free of organic matrix
and permineralization except for a calcitic crust found on some bones.
The bones collected by Zamyatnin and Kiladze were identified by
Gromova (1948) and later by the author. The list of species is given in
Table 13.
TABLE 13. Mammals from the vicinity of Mgvimevi*
Species Number of bones ПБ af
individuals
Сара Три” М дол me oo) Poe 2
Во Ce T'a CSpot, ain co cee, om, Abe eMC we, oe 1
Раша sspclaietay о gestae Томе: т GENES о. 1
Nes cre cis че ата аи... ПТИ. 1
раме <cawad Tse оси... MMe ale alte 3
Ba Sto MT USC US aie: 2 oc ce MM acl ре co ola 4
Capnewer, CaUiGNSiC A ae. deer. се. Вы.
O\W iS: ебало Die, WYRE ss aes, aM athe. oe 3
Caprovinae (not identified more precisely )
Тотал их. 15
* The numbers include material received by the ZIN osteological laboratory after the 1948
publication,
The game animals in order of importance were bison, goat and ram, the
latter represented by a noteworthy species of large argali. Horses were
also abundant. The presence of cave bear and Panthera spelaea
indicates the age as Pleistocene, possibly Topmost Pleistocene. The lack
of deer, boar and gazelle specimens is due to the limited scope of the
excavation.
The Gvardzhilas cave is one ofthe Upper Paleolithic sites. It was
partly excavated in 1915-1916 by the engineer Krukovskii (1916), and in
1953 by Kalandadze and Tushabramishvili (1955).
The cave is located in the ravine of the Chirula rivulet, the right
tributary of the Kvirila, near the village of Rgani. The wide oval entrance
faces south; it is located on a small protruding platform nearly halfway
up the right wall of the canyon. Several grottos are located somewhat
higher on the opposite wall. The length of the cave is 30 m, the width at
the entrance is 19.5 m and the height approximately 4 m (Figures 54, 55).
According to Krukovskii, the calcitic ceiling of the cave collapsed three
or four times and two ancient cultural layers were buried beneath the fallen
rubble.
120 The tools from this cave are mostly cutting and scraping edges made
of pink chert. Many bone implements were also collected: points, darts,
awls (Figure 56). From the flint tools, the site was dated as Top
Paleolithic, Magdalenian-Azillian.
114
FIGURE 54. Gvardzhilas cave
Photograph by S.N, Zamyatnin, 1926
Krukovskii collected nearly 150 bone fragments and teeth of carnivores,
ungulates and rodents; 14 species have been identified by Smirnov Е
1924).
The collections made by Kalandadze and Tushabramishvili contain over
4,600 bones; the author's contains 78 bones.
All the bones of large animals are typical food remains of Paleolithic
hunters: broken epiphyses, diaphyses, skulls and teeth. A small number
of bones of insectivores, rodents and birds are derived from eagle owl
pellets. The bones appear very fresh with a yellowish beige color
1 and nearly intact organic matrix. Much of the enamel has retained its
natural color showing only an occasional rusty tint or darkened crack.
When the bones are scraped, they smell of dampness. Many show tooth
impressions offieldmice. The state of preservation shows that the bones
cannot be older than Top Pleistocene and they are more likely to be of
Lower Holocene age.
115
: 0 5m
9 Spring nary
FIGURE 55, Diagram of the Gvardzhilas cave
The list of identified species from all the collections is given in Table 14.
The collections from the two layers are combined for purposes of the count,
since there are no differences between their species compositions.
Nearly 900 fragments of diaphyses from Tushabramishvili's collection can
with difficulty be identified as large ungulates, bison and deer; of which not
less than 550 belong to wild goats, chamois and roe deer. In addition, the
collection contains 2,970 unidentifiable fragments of small ungulates and
carnivores (Figure 57).
The sizable collection of Gvardzhilas cave material clearly indicates
a considerable transformation of the ecological composition of regional
mammals which occurred there [in the Colchis] at the Pleistocene-Holocene
boundary. Horse, sheep and, probably, porcupine vanished from the
plateau and, of the southern steppe-plateau forms, only the Asia Minor
hamster remained. Since there are no reasons to suppose that the
tastes and hunting habits of the primitive tribes changed at this time, it can
be assumed that the disappearance of these species resulted from the
afforestation of the highland.
Smirnov, Gromov (1948) and Pidoplichko (1951) regarded the Gvardzhilas
cave mammals as а Stratigraphically and ecologically uniform assemblage.
These authors attributed the downward shift of the altitudinal zones to the
advancing glaciers of the Greater Caucasus. They cited as support the
fact that primitive hunters did not hunt goats and chamois further than a
day's march away with aheavy load, i.e., 15-20km. Andthey also considered
that Prometheomys schaposchnikovi Sat. (a present inhabitant)
of alpine meadows) could not have existed in the cave environs in a zone
of xerophilous vegetation such as exists there today.
116
122
Our observations indicate faulty reasoning on their part. The distribution
of game animals, in particular that of Caucasian goat, was wider than at
present. And even now, suitable ground for wild goat is to be found in rocks
6 km from the cave along the upper Chirula. In April 1954, the author
observed one eagle owl which regularly flew each evening from its day
shelter below the Gvardzhilas cave to the nearest subalpine meadows and
the forests of the Rachin ridge. This owl probably preyed on hamsters in
the dry cave environment and hunted Prometheomys inthe subalpine
meadows and forests.
It is evident that a discussion of the Gvardzhilas cave assemblage need
not consider the shift of zones nor the mosaic structure of the landscape;
neither should it preclude the possibility of two ecologically different types
of animals coexisting in the same landscape zone: the alpine and
mesophilous Caucasian goat and Prometheomys and the xerophilous
Asia Minor hamster.
The finds of Prometheomys and hamster remains indicate a
Pleistocene faunistic connection between the western part of the Greater
Caucasus and the Lesser Caucasus along the Imeretian Plateau and the
western Slope of the Surami Range.
TABLE 14, Mammals from the Gvardzhilas cave
Number of Number of
Species :
bones individuals
ара Ait CAWCES ICR WSS wads В ео ob do 0 a6
BrimaCSws Ail, CHROPASCHSS oa5600000000n000 6 oc Ou
МЕТ ре: УП ДЕЯ ее м fore fou Wile ео о
О ео бое ое sas
Ursus aretos meridionalis
О BERCLOS CAMCASICISS 646 5 o.a0 goo bib adogeo 6
о Sie EMSS вме ENB: повтор
GSMO. ИО, ль ob, Geo ооо ао ANG ICN Иво овен
Mustela nivalis caucasica
MCSOCEI COURS Eiki, Па Ch cisononovaoo00 nied
Prometheomys aff. schaposchnikovi
В С TOMAS еее Meee etic cians
Capreolus capreolus
СИИ CIA DIS Gioia оо ооо додевойа CHOPE co Chena ones leans
Capra cf, caucasica
NUPUEAPA ириса DA oo Sos nooo boo o oma od dic
Bison bonasus (caucasicus?)
eee ee соо ооо
I= 5— (op) @м ею Cr T= 59 fe
Ce
Ce оо
Ov i bo) Co) CoRR NSE NE bb)
ray
65
ооо оо уно ооо ооо не
oo Н*
о ооо ооо ово
оао ацово ооо sO) O80 ооо
The Upper Pleistocene faunal complex of northeastern Colchis is known
by the remains of game animals and eagle owl victims from five cave sites.
The complex has some features which are like and some which are unlike
the faunal complex of the Black Sea coast. The similarities are in the
occurrences of the mountain-forest ungulates and carnivores: wolf, fox,
117
123
bear, European wildcat, marten, boar, roe deer, deer, elk and bison, and
in the complete absence of the indicator species of the Eurasian plains, i.e.,
mammoth and primitive bull.
As opposed to the Black Sea faunal complex, a number of migrants from
the high steppes of southwest Asia are present in the Colchis complex.
These include the Asia Minor hamster, porcupine, argali, horse and
Equus hidruntinus.
These species migrated from the south into western Transcaucasia.
Their migration was facilitated by the presence of a karst relief (i.e., dry
soil) and, toalesser degree, by the arid climatic phase which occurred
some time in the Middle Pleistocene.
27227777
WD
LL
ti Wig
ey
77
|}
LiL,
TLL Rp
UY
KM iy
MWY)
My
(1
SARA AAA
у
ee)
4
FIGURE 56, Flint and bone tools from the Gvardzhilas cave (from Zamyatnin, 1957)
Forest animals (boar, elk, roe deer) were abundant in the complex,
occurring mainly on the marsh plain of Colchis. The European beaver
was an ancient relict.
The alpine species of the highlands were much more widely distributed
than in present time.
The Kodor and Tsebel'da ravines formed "ecological barriers'' for some
northern (European) species and for the highland species of southwest Asia.
These two faunas converged in Upper Pleistocene time in the region of the
barrier.
118
125
Thirty species of mammals of Upper Pleistocene age are known from
Paleolithic sites and other localities in western Transcaucasia. The species
are distributed as follows:
ИСИ GS ab lola Зое ов ое 2 Бозе АА О В ie enter sree 7
Chino preralNed: a о ee eh aac) Meee 1 Rerissodactylaliige. ее Aya 3
SATIN OLA wept ый ak 13 АОИ Sy д op On bn Oa oo ao 13
ооо Бе Gyo ооо ole d bo 1
It is Significant that mammoth, Bos primigenius and northern
forms (e. Go, вещшавев або wep, ое hare) are missing from the complex.
And yet there are no warm-climate southern forms in the complex either.
The main difference between the western Transcaucasian and the western
Ciscaucasian complexes is the presence in the former of forest and
mountain species. This indicates that the contemporary fauna and landforms
were developed in western Transcaucasia by Middle Pleistocene time.
The transition to the Holocene (postglacial) fauna in western
Transcaucasia was probably more abrupt than the changes which occurred
during the Middle and Upper Pleistocene. At the end of the Upper Pleistocene
the following changes took place in the fauna: the extinction of cave bear,
cave cat and cave hyena; the southern retreat of southwest Asian upland
xerophilous species (hamster, porcupine, argali); and a decrease among
local forest and mountain species (bison, goat, chamois, roe deer).
However, the fauna which has survived into the present has preserved the
features of a forest fauna.
The climatic change in western Transcaucasia since the Middle
Pleistocene probably evolved in different ways than in other regions because
of the montane character of the local topography and of regional climatic
peculiarities.
The paleogeographic picture we now have will undoubtedly be augmented
by further studies of the karst in Imeretia, Abkhazia and the Krasnodar
Territory,
A special search for Lower and Upper Pleistocene cave sites must also
be undertaken.
Central Transcaucasia
The regions of the Surami water divide, the upper Rion, Bol'shaya
Liakhva and the Gori depression are of great interest to paleontologists and
zoogeographers as the probable migration route of Neogene and Quaternary
mammals from southwest Asia. Pleistocene mammals occur in Paleolithic
caves and in diluvial deposits.
In 1952 Lyubin (1953) was the first to find Lower Paleolithic tools in the
Bol'shaya Liakhva valley and at Metekhskaya Prona near Dzhidzhoita, Morgo
and Lashe-Balta, at an altitude of nearly 1,500 m. The tools from Lashe-
Balta (hand axes, flakes, cores) are made of andesite and other quartzitic
rocks. АП the specimens had been washed out of a high terrace (probably
the 60 m terrace) and rounded and redeposited by streams. As in Abkhazia,
the bones, whichwere probably originally deposited in the terraces together
with the tools, were not preserved. The value of the paleontological material
from the lower layers of the caves is therefore all the greater.
119
FIGURE 57, Mammal remains from the Gvadzhilas cave
1 — jaw of (x2) Talpa caucasica; 2—jaw of Gulo ай, gulo; 3 — femur (X 2) of
Mesocricetus auratus; 4—jaw of Prometheomys schaposchnikovi; 5—
jaw of Cervus elaphus; 6-8 — М, astragalus and metacarpal of Capra caucasica;
9-10 — М» апа first phalanx of Bison bonasus
120
126
Caves of Mount Chasovali-Khokh, Paleolithic and younger beds
in caves of the Kudaro region in South Ossetia, were discovered by Lyubin
in 1955. The four most important caves were located in the right wall of
the Dzhodzhora rivulet ravine above the villages of Kvaisi and Chasovali.
The caves are in the dolomite, 1,700 m above sea level and 300 m above
the riverbed (Figure 58). Much material was collected from the upper cave,
Kudaro I, which is dry and slopes gently to the north from two entrances
facing south and east (Figures 59, 60). The excavations in 1955, 1956 and
1957 exposed clay beds, 2.5 m thick, with blocks of limestone. Iron-,
Bronze-, Copper- and Neolithic-Agetools and pottery occur in the upper
part of the beds. Middle and Lower Paleolithic tools occur in the lower
part of the section. The flakes, hand axes and points are made of quartzite
and are in part Mousterian and in part Acheulean. АП the beds contain
3 Kudaro ТУ.
Kudaro 1-1
FIGURE 58. Mount Chasovali-Khokh (caves indicated by arrows)
Photograph by author, 1957
bones brought into the cave by men and owls. The Lower Paleolithic bones
are mostly devoid of organic matter; they are light brown in color, and have
dendritic surfaces. Some bones were rounded like pebbles in a smallstream
which flowed through the cave in wet seasons. Fragments of limb bones,
teeth, phalanges and metapodia of carnivores and ungulates predominate.
The material is difficult to identify to the species level.
Upper Paleolithic tools and well-preserved bones were found at a depth
of 3 m in excavating the clayey-lime beds in front of the entrance to the
Kudaro III cave. Table 15 and Figure 61 show the many mammalian
species which have been identified from these bones; up to 1958, 40
identifications from the Pleistocene had been made.
121
Collections from the Paleolithic beds of Kudaro I include: 42 bones of
birds, 3 bones of lizards (Lacerta sp.), 43 bones of toads (Bufo sp.),
22 bones of frogs (Rana sp.) and over 30,000 bones of fishes, primarily
of Salmonidae. *
The abundance of fish bones in the Lower Paleolithic beds might indicate
that the beds were deposited under water, possibly in the paleo-Dzhodzhora
River, which might have carried the bodies of animals and abraded the
bones.
127 TABLE 15, Paleolithic mammals from Kudaro caves
Kudaro I Kudaro III
Lower and Middle | Kudaro II Upper
Paleolithic Paleolithic
Тара Spite ons folate tone a ane сы: 8 1 —
GCL TS SAE oo oles ео 1 =
Chiroptera, not determined below the generic level 27 —
Масаеа р в ооо о 2 =
Sans: Сиро оо... 27 10
GHG "SP fe ое а.о. Prt cee 3 =
Wail pie Sach. ariel Die Se ee ee: Ре oA ame oes 33 1
Grocwtayspelaea tr с ны 1 =
Wrstismsp ella evises 22. cats Bs ieee rte : ‚ 2,979 97
Мате сабота, о he oS 9 =
Misiteliasich nmasviall isi 22 al spauas ceo cde? wre ay at ae в =
Melles ев железа. О Е 11 ==
Са о бе ооо Не oe 11 =
PAMENE Ta реТаеа р: о пон, ев Aye 3 9
Pepin Gast ие stave т а 9 1
Carnivora, not determined below the generic level 18 —
Lepuis’ еб eumo\p ен иен. au. clea a aie Nats 12 =
а оао Ay ao opt ala 0 J.2-c cle b vag С 10 aa
Hivis Dix Bers емонта ен Sahel one Е 18 =
Al Vatetagarieh twit ll ia msi. ое На 2 a
Mesocricetus, ай ama tus сес = devia А 56 =
Мистогиз мари Tra. ор A esataol. eee cl ome ae 16 =
Mi atten SNUG cee csc Зы NID eae tS - 5 —
Prometheomys aff. schaposchnikovi .... 13 =
Muridae, not determined below the generic level 52 —
RNTNOWEMOSHED: sie: ов, О а cae st 2c bs о 6 т
ор С ое ое сор и. nile 2 =
Cervus chy ella phiuste, .. смс и. . 213 20
Cap reo Misi а са pre 0 5... 36 =
CApTa ©, СЕНТ И, ео 55 11
Теа ОТВ М eter en И, МНН 26 11
Ovis or Capra, not determined below the generic
LEVEL) CELTS AEE: Bete atte Stine Rei HO ree 189 —
Виолетта. дааа = 5 —
Small bone fragments of ungulates and carnivores —
Artiodactyla-Gamivoras 5 ao dit лан 13,013 135
* Ina communication on the 1955 excavation (Vereshchagin, 1957b) the intermaxillary bone of a salmon
was erroneously identified as a bone of a large lizard,
122
(128)
FIGURE 59, Diagram of Kudaro I
1, 2 —excavations of 1955-1958
As a whole, the Kudaro I fauna is indicative of dry plateaus and includes
widely distributed forms (wolf, fox, cave bear, badger), Caucasian mountain
forms (panther, marmot), and southwest Asian forms (hamster, sheep,
porcupine). The indications of the migration of southwest Asian upland-steppe
forms to the Caucasian mountains and of the survival of apes through the
125
end of the Pliocene are matters of particular interest. Porcupines,
sheep, a few goats and chamois indicate that the reliefwas quite uniform
and the climate warm and dry.
128 Comparison of this site with the Upper Paleolithic sites of Imeretia
suggests that mountain goat and other mountain forms existed in isolated
areas of Caucasia which were undergoing differential uplift during the
Pleistocene. In the Lower Paleolithic the upper reaches of the Rion were
uplifted as much as 500 m.
The Upper Pleistocene fauna of central Transcaucasia is represented
by single mammoth teeth. One of the teethof Elephas primigenius
from the Dusheti area has been described by Burchak-Abramovich (1946).
The tooth of a small mammoth was also found in the town of Gori in the
first terrace of the Kura River; according to Gabuniya (1952a), it belongs
to an Upper Pleistocene or Lower Holocene mammoth. It is probably of
the same age as the late mammoths of the Pyatigor'e area.
The presence of mammoths in the Upper Pleistocene of Transcaucasia
is not proof of a migration of ''northern'' forms to the south, but rather
is probably indicative of an endemic evolution of the proboscideans, such as
occurred in the western Mediterranean.
'129)
FIGURE 60, Excavations at southern entrance to Kudaro I
Photograph by author, 1956
1704 124
129
131
Eastern Transcaucasia
In post-Tertiary time eastern Transcaucasia was plain country
surrounded by mountains in the north, west and south, andthe sea inthe east.
During the Pleistocene the seas transgressed at least three times over
the eastern part of the plain, considerably reducing the living space of the
land animals.
The known localities of Quaternary mammals are on the marine terraces
of the Apsheron Peninsula and on the foothill river terraces in the western
part of the plain. Bone accumulations in caves are almost unknown in
eastern Transcaucasia. '
Lower Pleistocene mammals occur in littoral sediments of the Baku sea
on the Apsheron Peninsula. Skull fragments of a young rhinoceros
(Rhinoceros cf. mercki), large horse (Equus sp.) and hyena
(Crocuta sp.) were collected at a depth of 2 м incoarse, shelly
limestone with Didacna rudis Nal. and О. surachanica Andr. near
the Kishly railroad station, north of Baku. (The skull of the hyena was
previously identified as Hyaena striata (Bogachev, 1925a, 1938b),
which is impossible for the Lower Pleistocene.) A fragment of mandible
and a tooth of Elasmotherium from the 12-m level in No. 19 oil well
near the village of Binagady are in the PIN collections. This materialcomes
from an ancient Caspian terrace with Didacna surachanica Andr.
The southern part of the terrace underlies kir* beds which contain
the Binagady faunal complex (discussed below).
A lower jaw of a colt (Bogachev, 1938c), identified by Gromova (1949)
as Equus aff. sussenbornensis, was collected in oil-bearing
gravels of the ancient Caspian terrace which overlies the oil-bearing beds
at Khurdalan on the Apsheron Peninsula.
Fossil mammals are more common in Middle and Upper Pleistocene beds
than in the Lower Pleistocene. However, their dating is sometimes
difficult because of the poor state of stratigraphic knowledge and the
discontinuities of the Caspian terraces. Most of the material comes from
the Apsheron Peninsula.
The preservation of material in this region was promoted by considerable
changes in the relief caused by tectonic, erosional and eolian activity. The
ancient, transgressing and regressing Caspian waters in some cases obliterated
and in others enhanced the work of the forces operating on the land.
Following the major transgressions of the Caspian, cones of mud
volcanos appeared оп the Apsheron Peninsula. Аз arule, all the deposition
sites of post-Baku fossils occur in oxidized oil shows, i.e., bitumens and
asphalts.
Many animals died in oil and liquid asphalt which preserved the bodies
in a perfect state. The bituminous beds, because of their plasticity,
protected the bones from erosion by water and wind.
In most cases, Pleistocene fossiliferous sites were formed through the
compound action of freshwater and oil or asphalt seeps on accumulations of
mammal remains.
The best known are the Binagady and Kir-Maku fossiliferous localities,
although fossil mammals also occur at other places, including:
а jaw ofa camel (Camelus sp.) found in a ditch at Surakhany; skulls and
horns of red deer (Cervus elaphus) collected in the bitumens on
Artem Island (Bogachev, 1924, 1932); maxilla of a horse of the ''Asian type"!
* Kir, or bitumen, was and still is used in covering and sealing flat roofs.
125
(130)
р ПИТТ
OE ив
ПОЙ
ee 7 | $,
р Пт
\ |,
FIGURE 61, Vertebrates from the Paleolithic beds of Kudaro I
1 — intermaxillary (x 2) of salmon Salmo spp.; 2 — МЗ (х2) оЕ Macaca sp.; 3-5 — milk canine (x2),
M,and metatarsal of Ursus spelaeus; 6—upper jaw of Panthera pardus; 7—jaw of Marmota
sp.; 8 — дам (Х 2)of Mesocricetus aff, auratus; 9 —side and worn surface of tooth (x 2) of
Hystrix зр.; 10—phalanx of Rhinoceros зр.; 11—hornof Capreolus cf, capreolus;
12 — first phalanx of Cervus cf, elaphus; 13-15—metacarpus, first phalanx and astragalus of
Ovis cf, ammon
126
(Equus cf. caballus) found in the clayey breccia of Mount Bog-Boga in
Balakhany, 15 km northeast of Baku; bones of a horse known from the ancient
Caspian terrace on the Tertiary hills near the village of Ali-Bairamly;
remains of one Bos, recorded from the Baba-Zanan Tertiary hills near
Sal'yany (Bogachev, 1938c).
Binagady
The Binagady fossils are important in the study of the Quaternary fauna
of the Caucasus. The occurrences at that locality of plants, mollusks,
insects, reptiles, birds and animals of all ecological and morphological
types allows a detailed reconstruction of the ancient Apsheron landscape.
The locality was discovered in 1938 by a student, Mastan-Zade, who
was studying the Apsheron bitumens. The first excavations were organized
by Bogachev in 1938 (Azerbaidzhan branch of the Academy of Sciences of
the U.S.S.R.). The excavations continued until 1941 with the participation
of Kasabova and Sultanov and were resumed in 1946 under the supervision
of Burchak-Abramovich.
The primitive mining of kir over many centuries left holes and
pits of various sizes in the bone-bearing area (Figure 62) which
seriously interfered with the excavations.
In order to obtain a clear stratigraphic picture the locality must be
excavated in a series of mutually perpendicular trenches.
(132)
FIGURE 62, Binagady hollow remaining from bitumen-mining in area where rhinoceros bones were found
Photograph by author, 1941
127
132
133
The Binagady locality is on the crest of а hill 0.5 km southeast of the
village of Binagady, and 7 km north of Baku. The coast at its closest is
10 km to the south and 25 km to the north.
The bone-bearing area comprises approximately 12 hectares and is
located on a hilltop near the Kyrrar hill. The area is 54-57 m above
present sea level, and 48 m above the level of Lake Beyuk-Shor. An ancient
mud volcano (Kichik-Dag) lies north of the fossiliferous area; further
to the north is the meridionally elongated, saline Lake Masazyrskoe
(Mirdalyabi) and to the northeast, Lake Binagady. To the east is the saline
depression Kariatakh-Shor, beyond which rises the Balakhany Plateau.
Extending from the Binagady hill are oil-bearing salinas and the
saline Lake Beyuk-Shor, which stretches far to the southeast (Figure 63).
According to the Volarovich-Lednev map (1913-1929) and Bogachev's
description (1940b), the base of the Binagady hill is a broad, anticlinal
fold made up of Maikop (Oligocene) and diatomaceous (Miocene) clays and
of productive sandstones (Middle Pliocene). South of the hill is the Middle
Quaternary Caspian terrace. An eroded cone of the mud volcano Beyuk-
Dag, 105 m above sea level, is of Pleistocene age. The volcano probably
formed on the northeastern part of the Miocene fold in two stages. The
loessial sands at its base are dated Q,''(Riss-Wiirm) and at its top Q2>(Early
Wiirm).
The crest of the fold was eroded by waves of the transgressing sea.
As a result of erosion the oil from the productive beds seeped into the
coquinas of the ancient Caspian terrace and into the later Quaternary beds.
The pressure within the productive beds resulted in oil and asphalt seeps
at the surface of the fold and subsequent cementation of the overlying
sediments by asphalt. Consequently, a series of fossiliferous asphalt
layersof varying ages appearsin the syncline. The most recent asphalt
seep occurs on the southeastern slope of the hill; it probably formed during
oil formation in the last two centuries.
R= La FIG Lake Mirdalyabi Gif HW =
= : 7, И MONA ПОР ==
2 Wael. Gy GS,
Fry, Sta’ Mt. Kichik-Dag == \
ТАМ: Beyuk-Dag к АЗ ЧЕ Ze. == G@
WAYNE 5 ie i wee : ИЕ АЕ EA
| Whee Wht, IS, 9222222.
MIN is Be
и
НИ
РЕ
№
ft fet} АЕ y Ma tf)
Se anne к. AS : Lm
KC 1:42000
EA
FIGURE 63, Present topography of the bone-bearing localities at Binagady and Kir-Maku
The bituminous area extends to the ancient Caspian ''Khasar'' terrace
in the south. The following fossils are characteristic of this terrace:
Dreissensia polymorpha Pall., Didacna surachanica Andr.,
128
О. kovalevskii V. Bog., Neritina pallasii Lindh. It is possible
that the lower part of the bone-bearing beds correlates with the near-shore
coquina of the Khazar terrace. The terrace slopes gently downward to the
south, an indication of tectonic movements in post-Khazar time. It does
not extend beyond the northern slope, which, in Khazar time, was probably
dry land which was later subjected to strong erosion. If, as agreed upon
by many geologists, the landscape of the area at the time when the bone-
bearing beds were being formed was similar to the Recent landscape,
it follows that during the marine transgressions the area became an
archipelago of small islands and peninsulas.
The present-day relief of the Apsheron Peninsula was formed through
the interaction of tectonic forces and eolian and subaqueous erosion
(M. Mirchink, 1934). The incidence of shallow, elongated salt lakes in the
area is a result of wind erosion. Even the deep Yasamal'skaya valley,
extending from Binagady to the sea, was probably formed by wind erosion.
The Apsheron limestones which cover the Tertiary loams in places were
evidently not sufficient to protect the loams from erosion by northerly winds.
The Upper Pleistocene and Holocene produced tectonic and epeirogenic
movements which were responsible for much important change in the relief
of the Apsheron Peninsula.
134 According to Shlepnev (1947), the epeirogenic movements in the
Apsheron over a mere 16-year period (1912-1928) resulted in differences
in elevation between various points as great as 114 cm. Thus the present-
day relief in the Binagady area cannot be used as a starting point in the
restoration of the relief at the time of bone deposition.
A generalized stratigraphic section from the center of the bone-bearing
area is given in Figure 64. Surface layer А is diluvium, 0.2 to 1.0 м thick,
underlain by bedB, which is 0.75 to 1.4 m thick and contains fine-grained
sand with fairly coarse coquina of marine mollusks, impregnated with oil.
The southern portion contains large trunks and branches of juniper and
occasional bones of horses, deer and large birds. BedC consists of dense,
viscous, dark brown bitumen, 0.1 to 0.5 m thick, which seems to form a
lens toward the margins of the bone-bearing area. Pockets near the top
of the bitumen bed contain bones of birds (mainly ducks), beetles and grasses.
Bones of large animals are embedded in the bitumen of bed C, and
occasionally at the boundary of beds B and C.
BedD is fine-grained sand with fine coquina and some small pebbles;
it measures 0.3 to 0.6 та in thickness. Horizontal bedding, produced by
streams, and films of narrow, swamp-plant leaves occur in this bed, which
is the main bone-bearing bed. It contains bones of mammals, birds and
reptiles, as well as insects, mollusks and plants.
The granulometric analysis of beds BandC shows that the fine-grained
material amounts to 62.5 and 75.4%. This probably indicates the littoral
eolian origin of the Binagady beds (Vereshchagin, 1951b).
Fossil animals and plants occur in pockets in the oil-impregnated sand
which underlies the bitumen bed. These pockets are more heavily
impregnated with bitumen than the surrounding rock.
The osseous strata 3 m from the surface gradually grades into the
fine-grained sands of the productive beds, which are slightly impregnated
with oil. A bitumen bed, 10-15 cm thick, can be seen in the holes dug at
135 the highest point of the hill. This bed, 0.8-0.1 [?0.8-1.0] m from the
129
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ЕЕ — М = М — И=/л
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ИЕ ИИ —Й
| at
0.2-1.0
07-14
lle
03-05
surface in Holocene loams, was probably
formed during a recent phase in the
migration of oil. It is rich in bones of small
birds.
The origin of deposition of the beds shown
in the section is not entirely clear. Bed B
was probably formed along the shore ofa
body of standing water, as indicated by the
coarse coquina of marine shells composed
of fragments 2-5 m in diameter. Bed C was
probably formed as a result of migration of
oil along a fault in the productive beds
somewhat to the north of the Pleistocene
bituminous lense.
Mastan-Zade (1939) has hypothesized
that an oil flow occurred during the eruption
of the Kichik-Dag volcano.
It is obvious that bituminization (oxidation
and thickening of oil) occurred at the surface.
This being the case, bedB must have been
formed during a transgression in the basin
which covered the bituminous lens, since
the bituminization could not have occurred
under water or else the oil floating to the
surface would have been dispersed by wind.
Bed D was probably formed in a short-
lived body of stagnant water. It is difficult
to detect the boundary between the osseous
bed and the productive sands, because it
is not clearly defined. The negligible
percentage (0.2-1%) of shell fragments,
2.0-3.5 mm in diameter, indicates littoral
marine or estuarine deposits, in addition
to windblown, fine-grained sand.
The incidence of xerophilous grasses,
small tree branches and detritus mixed with
animal bones and fossil beetles indicates
that the material was deposited in topographic
depressions from small mud-oil flows.
Individual bird bones, remains of insects
and plants, skulls of animals filled with bituminous material — all these
indicate poor sorting of the kind that occurs ш а river flow. The likelihood
is that the remains accumulated in a stagnant basin where the bones were only
slightly moved over the substrate.
With the exception of individual skeletons of small birds (dove-size or
smaller) all the skeletons at the site are disjointed. Remains of ligaments
and feathers occur only on the bones of birds from the younger beds, e.g.,
the bituminous intercalation in bedB. No complete skeletons of large
animals were found in the first excavation, possibly due to the scale of the
excavation. Scattered bones of large and medium-sized animals occur in
the asphalt beds mostly in horizontal positions. Burchak-Abramovich has
found bones of a horse and of a deer scattered over an area of several
FIGURE 64. Section through the asphalt
beds at Binagady
А — surface loams; В — bituminous sands
with coquina, bones and wood; C— bed
of "rich" asphalt; D— oil-impregnated
sands with animal and plant remains;
E — sands of productive beds. On the
right — thickness in meters
130
square meters. In the bituminous bed complete skulls of medium-sized
animals (wolf, hyena, badger) were found separately from the cervical
vertebrae; even the atlas was separated from the skull. One find has been
recorded which consisted of six joined cervical vertebrae of an ass and
the mandible of a horse in two sections: the anterior part, which had been
broken off, was deposited in a bed adjoining the horizontal layer where
the posterior section lay. Broken bones of large animals and birds occur,
as arule, inthe lower part of bed В.
The deposits in bedD are characterized by quite complete bones,
136 including some broken epiphyses and intact skeletal bones of large birds —
all with asphalt-filled cavities as a result of the penetration of oil into the
bone tissue pores.
One seldom finds bones that show tooth impressions of wolves, bears
or hyenas, although gnawed humeri of bears, the pelvis of Equus
hemionus andthe femur of a rhinoceros have been described by
Vereshchagin (1951b) and by Burchak-Abramovich (1953a). There are no
signs on the rodent skulls of beak or claw puncture by birds of prey, which
indicates that the bitumen-preserved skulls probably were not deposited
in the excreta or stomachs of birds which fell into the asphalt. Puparia of
flesh flies occur, although not frequently, in nasal and brain cavities, and
in the depressions of a rhinoceros humerus. When the puparia were opened,
it could be observed that the flies had hatched from the pupae. There were
no bones found which displayed cuts or breaks made by prehistoric man.
Paleopathological evidence of osteomyelitis, rachitis and helminthiosis
is common (Vereshchagin, 1946b, 19515; G.V. and D.V. Gadzhiev,
1952). Only two bones of a wolf and deer with polished distal epiphyses
are known. Among the Binagady bones, there are only very few which have
been rounded by streams, or which show sand-polished surfaces or other
signs, such as brittleness or cracking, of origins in land surfaces or
well-aerated soils.
The extent of permineralization of the bones is relatively slight. The
diaphyses, when they are sawed, smell of damp bone, which is unusual for
Middle Pleistocene material. Proteins were detected in the humerus ofa
wolf, and juniper wood from bed В contained 30.52% cellulose (Mamedaliev
and Kaplan, 1948).
The dry bones and polished surfaces of diaphyses are dark brown and,
in some places, bronze. In fresh fracture the bones show a grayish brown
or chestnut color. Upon being boiled in alkali, the bones become brittle
and light chestnut in color. The enamel and dentine of teeth are usually
gray-brown to chestnut. However, the large molars of hyenas, saigas and
bulls are bluish in color on the inner surface.
Because the extensive excavations of 1938-1942 at Binagady were poorly
documented, very little is known about the nature of the deposition.
Almost all the investigators who worked with the Binagady material were
concerned with the Apsheron landscape and the origins of the deposit: the
reasons for death and the circumstances under whichthe remains accumulated.
Bogachev (1939c, 1940b, 1944) maintains that the present-day relief
is very old and that the contemporary Lake Beyuk-Shor is a relict of an
ancient and much larger lake. He thought that rhinoceroces, horses, Bos
and deer attempting to reach clear water were trapped in the oil-surfaced
lake, and that quadruped and feathered predators attracted to the floating
bodies fought and died over their prey. According to Bogachev, the region
131
137
138
was also inhabited by migratory birds, and numerous streams
presumably entered the lake carrying plant detritus, juniper trunks and
pear fruit. He assumes the climate of the region to be somewhat colder
and more humid than that of today.
Considering all the evidenee, Bogachev compares the locality with the
asphalt deposits at Starina (Galicia) and at La Brea (Los Angeles, California).
He points out that organic remains accumulated gradually without any of
those catastrophic episodes which kill thousands of animals of
morphologically and ecologically different species.
There are, however, a number offacts which donot fit into this elegant
picture. If the relief in the past were similar to the present, then the
Binagady hill would have been a small island when the level of the lake was
higher and there would not have been room for all the animals nor for the
accumulation of oil on such a small island.
The absence of freshwater mollusks and fish in the bone-bearing bed
and the concomitant incidence of xerophilous plants do not support a picture
of a large freshwater lake. And, indeed, if there had been numerous clear
streams, there seems little reason for animals to have waded into oily mud.
The idea of rhinoceroses and boars dying trapped in mud can practically
be dismissed. There are no silt deposits in the Binagady beds, and the
animals could not sink in the very thin layer of fine-grained sand and
coquina.
From the plants identified in the asphalt, the botanist Petrov (1939)
has inferred the existence of savannah and sparse arid forests in the
Apsheron. He thought the site of deposit was in the delta region.
According to Argiropulo and Bogachev (1939), the occurrences of forest
dormouse, red-tailed gerbil, migratory (gray) hamster and jerboa, and
of mesophilous species of beetles indicate that the forests were of tugai
type and the climate was more humid than now.
The ornithologist Serebrovskii (1948), in an analysis of the avifauna,
included migratory birds in the species composition. He visualized the
area of the locality as a shallow, reed-grown swamp Surrounding a marine
bay. Bird casualties were attributed by Serebrovskii to oil film on the
water.
According to the geologist Sultanov (1947), the deposit site was located
on the shores of a shallow marine bay beyond the main zone of wave action.
His conclusions that rhinoceroses, Ellobius, mole voles and jerboas
drowned in the sea and that flying birds died of fumarole gas poisoning
do not require comment.
Efremov's statement (1950) in his excellent ''Taphonomy"' that bone-
bearing bed D is alluvial in origin and was presumably protected by the
asphalt from later erosion is in error.
Gromov (1952), who studied the rodents, adopted Efremov's taphonomical
viewpoint and explained the accumulation of organic remains at Binagady
by the agency of river sediments. He assumed the existence of treeless
steppes on the Apsheron in the Upper Pleistocene.
All the authors mentioned above have failed to explain the occurrence in
one and the same bed of plants, mollusks, insects, birds and small rodents,
medium-sized carnivores and large, hoofed mammals. Even though the
data confirm the deltaic origin of the bed and the transport by river of
animal remains, other questions, such as the causes and places of death
and the distances traversed by the remains, have not been answered.
132
However, ап analysis of the available geological
and biological data can give a true picture of the
Binagady site at the time of animal deposition
and of the landscape of the Apsheron in the
Pleistocene.
In addition to Binagady, we studied cases of
mass death among insects, birds and animals on
the Apsheron and in Kabristan and the conditions
of continuous accumulation and burial of their
remains.
A study of geological sections in both older and
newer excavations showed that a lacustrine or
river origin of bone-bearing bedD is doubtful.
The osseous strata are always found beneath
asphalt bedC, and when C wedges out, the organic
remains disappear. This can only mean that the
oil-asphalt provided the prerequisite condition for
the accumulation of organic remains.
Observations in Kabristan near Khurdalan,
Kir-Kishlag and Beshbarmak show that
incrustations and accumulations of asphalt occur
oe on mountain slopes and in depressions, near
munis with closely spaced : | : а
с peculiar fissures from which the oil is forced to
bearing bed at Binagady the surface. Gas and salt water are ejected, in
addition to the asphalt, by the mud-oil craters
or wells. The margins of the latter are overgrown
by sparse reeds (also found in Binagady, Figure 65), the green of which,
in the spring, makes a remarkable contrast to the yellow background of
semidesert, mountains and salinas. The wells attract the cattle, and at
Beshbarmak cows and asses died after sinking into the thick mud of the well.
Rainwater often accumulates in the asphalt crusts. When the asphalt
is hot and soft, the water forms lenses which are fatally attractive to
animals in periods of drought. Asphaltic pools such as these annually trap
many animals, and remains of various sizes, from that of mouse to
that of goitered gazelle and ass, can be found in them. The decaying remains
sink into the asphalt andthe bones and mineral particles brought by winds and
streams become incorporated in the silty beds underlying the asphalt.
It is not essential that the asphalt pools be located in large reservoirs,
valleys or gulleys for animal remains to accumulate in the asphalt. On the
contrary, asphalt located on gentle slopes or even on the crest of a ridge
is guaranteed long and effective action, because on such terrain it is not
buried under sediments (Vereshchagin, 1951b).
These Recent examples indicate that animal death in the Pleistocene
might have occurred at or near the site of deposit, without necessarily
involving any transport over long distances.
Over the centuries, the sticky surfaces and oily freshwater pools created
by the heavy Binagady oil seeps on the shores of the marine bay probably
139 became poisonous traps and burials for a variety of animals (e.g., hares,
jerboas, gerbils, voles, hamsters) and for many types of birds and insects.
Heavier animals such as rhinoceroses, horses and bulls would have been
supported by thicker layers of asphalt. This herbivore carrion undoubtedly
FIGURE 65, Upper part of the
stem of Phragmites com-
133
attracted carnivores and scavenger birds which accounts for the many bones
of foxes, wolves and hyenas in this area as compared with their paucity
on the river shoals. The author has described elsewhere the pursuit by foxes
of birds which were trapped in the asphalt (Vereshchagin, 1946a).
Transport of bodies by water with decomposition occurring at the site
of deposit could have taken place only on a small scale.
The absence at the site of mesophilous vegetation of forest and meadow
animals and freshwater mollusks and fish would indicate that the streams
which carried the bodies must have been small and ephemeral. They were
probably short-lived torrents originating not more than a few kilometers or
more likely, a few hundred meters distant from the site.
Bed (with coquina and juniper trunks) was probably formed during a
marine transgression over a shallow estuary. It covered the bitumen and
contributed to the consolidation of the bone-bearing bed. During the erosion
of the southern part of the osseous lense, it would have been possible for
bone fragments to be redeposited and incorporated into bedB. Tree trunks
carried into the bay by waves were also preserved in bedB (Figure 64).
TABLE 16, Age groups of wolves at the Binadagy locality
Number of specimens
Age
absolute 8
Approximately 1 month
20 3.5- 4 months
15 7-8 months
61 1-3 years
20 3-5 years, with slightly worn teeth
Over 5 years, with heavily worn teeth
Such biological data as the seasonal mortality pattern and the composition
of animal populations can contribute greatly to an understanding of the
nature of the Binagady locality. Clearly, the insects and pear fruits could
only have been trapped in the asphalt in the warm seasons of the year. The
incidence of migratory birds both from the south (spoonbill, purple heron,
black stork) and from the north (snowy owl) indicates that bird deaths
occurred during both the summer and winter seasons.
Death occurred at all ages, from infancy to senility, advanced age
being indicated by some individuals with molars completely worn down to
the base. Wolves are a particularly clear example. Because of the greater
size and strength of the wolf bones, it is easier to extract them from their
matrices than bones of other species, and thus to form estimates of the
size of the wolf population.
The age groups of wolves based on their mandibles are given in Table 16.
The numbers clearly show that animals of all ages, not only the very
young and very old, were buried in the Binagady beds. Data in Table 16
indicate that the main part (67.4%) of the fossil wolves are adults
(Figure 66). In other words, the dead assemblage represents not a selected
134
140
142
FIGURE 66. Mandibles of wolves of varying age
groups from Binagady
(Illustrates Table 16)
age group, but a cross-section of the
animal population which lived in the
area throughout most of the year.
Analysis of other carnivore fossils
also indicates that the animals
perished throughout the warm season
of the year (Vereshchagin, 1951b).
Taken together, these facts show
that the Binagady locality itself
provided the single cause of death at
this site, rather than at some
other locality.
From these preliminary
assumptions it is possible to evaluate
with greater certainty the faunal
complex at Binagady, as well as the
ecology of that period in eastern
Transcaucasia.
According to calculations by
Burchak-Abramovich and Dzhafarov
(1953), nearly 8,213 bones of at
least 412 individual ungulates were
collected during the 1938-1942
excavations. Carnivores are
represented by 2,043 bones of 11
species and at least 440 individuals,
lagomorphs and rodents by
approximately 2,300 bones of
15 species and at least 180
individuals, andinsectivores by nearly
50 bones of three species.
Bird bones total at least 30,000.
Hundreds of bones of turtles, lizards
and snakes, many thousands of insect
remains (mostly beetles), and
hundreds of land mollusk shells have
been found. Thirty-nine species of
mammals have been identified (Boga-
chev, 1938b, 1940b; Argircpulo, 1941b;
Vereshchagin, 1948, 1949с, 1951b;
Burchak-Abramovich, 1951c, 1952а;
Dzhafarov, 1950, 1955; Alekperova,
1952, 1955; Г. Gromov, 1952, and others).
We estimated the number of
common forms from individual
mandibles andthe number of rare species from all the bones. The statistics
on ungulates were taken from Burchak-Abramovich and Dzhafarov (1953).
The data in Table 17 is taken from collections which, in 1945, were in the
museums of Baku, Leningrad and Moscow. The quantities shown for bones
and individuals reflect not only the actual number of animals, but also the
method of extraction from the rock matrix. For small species, the
quantities shown are less than the actual numbers.
135
TABLE 17, Mammalian species and number of individuals from the Binagady locality
(141)
дв Number of в Number of
P individuals р individuals
Insectivora Mesocricetus auratus plani-
3
Croiedidiirak або т wisis ayes ss res 3 : .
; Е Meriones erythrourus inter-
Hemiechinus aff, auritus..... 3 :
Erinace aff, europaeus 1 ен =
u SBE MTG Pra Cae нь.
at : P Allactaga elater dzhafarovi 4
Sub tyo,t all eee vacate en oe 7 A. williamsi, 10
A, jaculus bogatschevi 26
Carnivora Dyromys nitedula 1
Mi t a li
Canis lupus apscheronicus.,, 120 ‘SoM Mae ee ee 70
: М. зоста 11$
Сами мере 56 т :
М. apsheronicus 2
Vulpes CONS AG ese в: 2 пара 85 :
i Ellobius aff, lutescens 10
VeVi реза: ато К oe coe 125 Е Е
Arvicola ef, terrestris 1
Е Е SR Г. Hystrix vinogradovi 25
Ursus arctos binagadensis ... 4 у Е
Уоттеаарежщея ие пана. пе ast: 13 Subtotal 177
M elesemeles! а: лишено 55
Рапенета spellers errs pri 11 Perissodactyla
ес Iybiician о. tlie 2 -
Ай г а | a Equus aff, hidruntinus 73
Yep peRt о SAE Ahi Е Е. caballus subsp, 154
Sib otal) р Se 440 Rhinoceros binagadensis.... 31
Subtotal 258
Lagomorpha
Териз Curopaeusigureeviors® .F. 4 Artiodactyla
ВЕ ce eles 4 Sus apscheronicus ae 11
Мезасевоз св. епусево $ в. 2
Rodentia Cervus elaphus binagadensis 52
Mus: шие снос 2 wee ол: cee BP
: Ovis cf, ammon Е 1
Apodenm's sylvatiens 7... . 3 6
Cricetulus migratorius
заримо pay Tio еее eas fhe 8 154
1,040
TABLE 18, Avifauna, by orders, from the Binagady locality
Number of
species
Number of
species
Limicola CunA0.) ga le LUGAR ae te NGoraciifornes \. соб a) cic trey: 6
ArisGhes ааа т. заза, ait Stegamopodes: 4. 5 лова ыь tate 45 2
AGCIPIIES 5) syesryen de clad) Pores og ь Гапона еь плз ть ь дла 2
GIEIROTES, 5 конь Вас Е" PIeTOCleleS ЕО о ой к РА 1
Е НЕ ее, О а ое а 1
Pee я Wie. dhe, РОН = 00. 7 as) > cua nen ets 1
136
Characteristic bones are shown in Figures 67, 68, 70 and 71.
The total number of bird species, as given by Serebrovskii (1948) and
Burchak- Abramovich (1951с), is 97 (Table 18).
Among the reptiles, bones of the grecian tortoise (Testudo graeca
L.) and small unidentified lizards are known.
Insect remains are abundant, but only beetles are well preserved.
A. Bogachev (1947) has described 107 species of beetles of the following
families:
Cicindelidae Dermestidae
Carabidae Hydrophilidae
Dytiscidae Coccinellidae
Cyprinidae Tenebrionidae
Staphylinidae Cerambycidae
Silphidae Chrysomelidae
Histeridae Curculionidae
Buprestidae Scarabaeidae
Kirichenko has identified six species of bugs of the Thyreocoridae,
Pentatomidae and Gerrididae families.
\ | | рота
FIGURE 67. Binagady
Mandibles: 1—Hemiechinus auritus; 2—Erinaceus europaeus
Remains of flies, fly puparia, butterflies, dragonflies and grasshoppers
are poorly preserved and difficult to identify. Shells of mollusks belong to
local land species of Helix, Helicella and Pupilla.
Plants identified by Petrov (1939) and the author (Vereshchagin, 1949c)
are represented by the following forms:
Pirus salicifolia Juniperus polycarpos
Pistacia cf. mutica Punica granatum
Vitis cf. silvestris Allium sp.
Zozimia sp. Phragmites communis
Tragacanthum sp, Colchicus sp.
Scirpus sp.
Bogachev (1940b) has also reported Alhagi, Isatis, Artemisia,
Salsola and Tamarix fromthe bone-bearingbed. However, these finds
have not been confirmed.
Some of the tree remains listed above come from coquina bed B, and
thus are younger than the main bone-bearing bed. This does not, however,
] 57
143 exclude the possibility of a stable xerophilous landscape in eastern
Transcaucasia.
The paleontological material from Binagady is so complete that one can
speak of entire Pleistocene complexes of different taxonomic groups and
biocenoses of Transcaucasia.
An ecological analysis of mammal assemblages reveals specific features
of the Apsheron landscape, climate and zoogeography. Changes which
have occurred since the Pleistocene in faunal species composition should
be projected to the level of orders (Tables 19, 20, 22, 24), taking into
account the Recent zoogeography of the northeastern and southeastern
periphery of the Bolshoi Range.
Insectivora
The absence of shrews and moles from the Pleistocene of Apsheron
indicates axerophilous landscape. At present, the long-tailed white -toothed
shrew inhabits forests and semidesert, andthe long-eared hedgehog the steppe
and semidesert, sometimes frequentingsparse, aridforests. The European
hedgehog lives in damp forests and in the dry bush of the semidesert.
TABLE 19, Comparison of Quaternary insectivore species in the piedmont of the eastern Caucasus
Pleistocene
Historical time
Species
Apsheron and
Kabristan
Dagestan
piedmont
ee eno eer ee) en OR On
Sieh вре ани 0% ey erie, ар бе о
о а eis te) cle) oe of os, 6, ete 5
о ООО
Note, The query (?) п this and similar tables indicates species likely to occur within the given period
of time.
Carnivora
The absence of forest species of carnivores (pine marten, European
wildcat, lynx) excludes the possibility of extensive forests on the Apsheron.
The lack of otter and mink indicates that there were no rivers or reed-
grown lakes rich in fish. Since weasels and stone martens are rare on
the Apsheron in present time, they may not have been preserved. The
presence of European brown bear (at present found in the sparse, arid
forests of Transcaucasia) suggests that the ancient Apsheron was a dry
plateau with juniper and pistachio forests growing along the creek beds.
138
144 The presence of small fox and corsac fox is further evidence of the resemblance
of the ancient landscape to the present. The incidence of cave bear is notable
and can probably be attributed to the dry climate.
Fast-moving carnivores were also common: wolf, fox, corsac fox, cave
hyena, badger, Panthera leo, cheetah, African wildcat. These animals
are adapted to the steppe or to brush and wooded areas of semidesert. The
climate of the region might have been colder than it is in the present.
However, the snow cover must have beenthin for the African wildcat, corsac
fox and fox to have been able to hunt.
In order to clarify the nature of the deposit and the evolution of species
assemblages from the Pleistocene to the Recent, the number of carnivores
at Binagady was compared with estimates of Recent carnivores based on
contemporary commercial game hunting in the Shemakha area near Baku.
The study revealed some similarities and some differences in the ecology
of the two assemblages, and changes in the proportions of five common
forms: fox, wolf, corsac fox, badger and jackal (Table 21).
A comparative examination of carnivore species of both periods confirms
the fact that the characteristically mesophilous Shemakha forest of the
Recent, which is the habitat of martens and cats, is not at all similar to
the Binagady trap and mountains.
Lagomorphs and rodents
The rodent species composition was almost twice as rich in the
Pleistocene (Table 22) as in the Recent on the Apsheron. Their ecological
grouping resembles that of the Recent Dagestan piedmont and indicates
that, in Pleistocene time, Apsheron was a piedmont steppe marked by small,
dry forests and some lakes which were overgrown by reeds and cattails.
Common hamsters live in an environment more mesophilous than the
one that exists at present on the Apsheron (see Chapter III). The presence
of the common and the Apsheron vole is another indication that the habitat
was more mesophilous than now. In contemporary distribution patterns,
the lower boundary of the common vole lies in the piedmont brush near
Shemakha at an altitude of 700 m, whereas the pine vole does not descend
below approximately 1,000 m.
Other species (hare, migratory hamster, steppe mouse, common
vole, red-tailed gerbil, jerboa, mole vole and porcupine) form an
assemblage characteristic of the present-day xerothermic valleys of
northern Iran and Karabakh, which are covered with spiny astragali,
willow-leaved pear, juniper and hawthorn.
Comparison of the Binagady rodents trapped in oil pools with those
caught on the Apsheron in the Recent by eagle owls and by personnel of the
Azerbaidzhan anti-plague station, indicates that in the Holocene the
proportion of xerothermic forms gradually increased at the expense of
mesophilous forms, which ultimately disappeared (Table 23).
Man is not responsible for the observed decrease of rodents from the
Pleistocene to the Recent (see Chapter VII).
139
145
FIGURE 68. Binagady
1,2—skull of Vulpes согзас; 3,4—skull of V. vulpes aff. alpherakyi; 5, 6 —skull of Canis
lupus apscheronicus; 7, 8 — canines of Ursus arctos binagadensis; 9-12 — skull, atlas and
humerus of Crocuta spelaea; 13-17 —skull and humerus of Meles meles aff, minor; 18 —
skull of Vormela peregusna; 19, 20 — jaws of young Acinonyx aff, jubalus; 21 — humerus
of Felis aff. lybica; 22, 23 — jaws of young and 24 — jaw of adult Panthera spelaea.
140
(148) TABLE 20. Changes in the species composition of the Quaternary carnivores in the piedmont of the eastern
Caucasus
Historical time
Specigs ны о апа Dagestan
Kabristan piedmont
Сами аси о р бое оно о, о о обв = + +
Саи ор оо ses) oh Е ОИ а ие Я + - -
Со MODES oc os oo ооо вов оборо о сво + + +
Уц [рез еда... . О юм... + = -
М. ми 1 резиу. ..... ПВ. ...... + + +
I WASmey SUTIATA Gog Gelb оо во вобововов = + +
Сто паре Таха see elect ees eon + - =
UWRGWS ВОГО особь И ооо + F oF
Мотитетамреже ло тиай. с. ogous aaas + a +
Miulsitie dl aaah У (gts mame о с = + +
№: ia tr ROM аще. Bile esses oo м... BA a = = +
Mi, GWEESMMAMIN Goo oecopogso00ndobacan = - +
ЕЕ ЕЕ сора бь юр ою оо во @asue = = +
Матее$ BRON Male Gi. оо... = + +
Меце пе Пе бесы uate ss lees sc Rees ves Ms Gs + +
PAS 2 ре аа... > GONG, o ome > =
IMINO оно модов р So ОН. < Ве = 2 ?
Pee pairiGiuteas, с. ос соо неее. = + +
SUNS ЛУ овобаво сво ооо о 0 Gago ов = + +
A SASS iit с Nese oscar chow Omoinaaeeenn once = В т
Ro ПУДРЫ оо о бо В о ое сои Stic + ? 2
Ее с Па А, о ce eee Ва ВН oe ee - + +
A CAIN i ух ЕВ о ББ боров об ооо or + ? ?
TABLE 21. Relative proportions of carnivores in Apsheron (asphalt trap) and in Shemakha area (commercial
game hunting data)
Apsheron in Pleistocene Shemakha area, 1930-1940
percentage percentage
species of species of
individuals individuals
ори: Ро о ео о Meath ФО о d oan tloo bg Ooo нояб ас 75.5
МОЕ о. c's а чет BENGKA MaMa Е 16.4
Gorsac оо Jackal ,„.... ое 3.0
Badger’, 20) . ое” es Wolf ес + Sennen : 2.3
Cave Эт Б во БМ eee о European wildcat and jungle cat .. 0.8
Tigempolecat а оао боев Pine marten and stone marten.... 0,7
INNO G30 co о боя об oto GO посоюо вс Beatie в ее обеих © 0.2
BEBE soe agalakvoondod ogo ово LYS ооо босоро ово особо сб 0.2
Е keys ebeiteoutin ale WAGE cobb oo A оо co ong oor 0.1
Я! ПИЩЕ ob Oo a oo ee oka OW bo eens 0400 осо ово 0.04
Мо! (59а): 2 ле bee оо эо с Taisen pole cai ев. Неее =) В ее нь ee 0.04
— ЗРЯ NOTE) Боба bod de 0% 0.001
= AVON о 6 Яо дао о моб оч 0.0001
Numbeniotispeciess uel. ce Number of species ..... 15
Number of individuals ... Number of individuals... 12,890
14]
146
FIGURE 69. Binagady
1,2—jaws of Lepus europaeus aureevi; 3-6—femur, metatarsal and tibia of Allactaga
elater; 7-10— jaw, femur, metatarsal and tibia of A. williamsi dzhafarovi; 11-15 —skull,
femur, metatarsal and tibia of А. jaculus bogatschevi;
142
(147)
FIGURE 69 (continued)
16, 17 — maxilla and femur of Cricetulus migratorius argiropuloi; 18-21 — maxilla,
femur and tibia of Mesocricetus auratus planicola; 22-26 — maxilla, humerus, femur
and tibia of Meriones erythrourus intermedius; 27-30 —skull, femur and tibia of Ellobius
aff, lutescens; 31-32 — maxilla and scapula of Hystrix vinogradovi
143
149
TABLE 22. Changes in the species composition of Quaternary lagomorphs and rodents in the piedmonts of
the eastern Caucasus
Historical times
Pleistocene
Apsheron
Species
Apsheron and
Kabristan
Dagestan
piedmont
Le pus ещораец$ В. . еее
Citellus рузшаев........ ce ees
MiCrOM у тов. еее.
Миз museugseteye - 2- « - а eee
А воен "sty livatt vest, see seen. he meme
А. ауте >. ров 2 Час
А. ag ta Wi WS «se © © > oo ео,
Cricetulus supgratorgus . . ieee ...
Mesocricetus auratus planicola..
М. auratus@mmpriculusr .... fue «oe
Meriones етот .... И > ..
А 1астази и acon ruan: .... м...
Allacta gia Че\атег. . Be. . a = + а...
A. williams. 2... Zaks сс...
А. JasCw Li See «cs с, с. ое
Dy tom ys iameaduliay, a. 2 Ме.
ОО elias SSS Fc eto aed au
IMIS ОНА 5.4 B Sitedicrd nodule 'g ol oa Oo
М. AD S:C ll COMME US) 5 деи газа
ElLobiws “aie ute semis wry ee. es. cow ими,
Je BEN opis WES, Бао ое ды О oon) с АО
АтутсоТа "HERE C's О, се
Hystrix утшорта ом, со, 5.
че
* Recent black hamster is a possible descendant of the Binagady species,
150 Perissodactyla
The presence of Equus hidruntinus, horse andrhinoceros (Table 24)
indicates that the Apsheron in Pleistocene time was a grassy steppe with
bush-grown gulleys. The grasses were probably burnt by midsummer.
Artiodactyla
The Pleistocene boar on the Apsheron could have lived in estuarine reeds
and in the pistachio-juniper forests. Boars now live throughout the year
in the thin, arid pistachio-juniper forests of Transcaucasia. The presence
of saiga indicates wide-open areas with steppe vegetation. The Binagady
Bos and deer might have entered the Apsheron during the spring growth
of grasses. The decrease of artiodactyls on the Apsheron near the beginning
of the historical epoch was caused by man.
144
(149) ТА BLE 23. Changes in the species composition and proportions (in %) of rodents and hares оп the
Apsheron from Pleistocene to Recent
Pleistocene, ИН || 1939. 1940 | 1947,
/ | centuries, ¢ :
Species caught in enna caught in caught in
asphalt pools Saye eres traps oil pools
Sipe WOME о co Bao ооо въ особововв 38.6 23.8 | 4.6 25.0
Comin) уе о но, она о. } i Absent
GICAE ETD OAM.) рес ARO nar bb 14,4 Absent
Втразаау DIOMOUPINE 5 Gace 500c 0b а 13.8 Extinet
Rec=iailesl Буа РОЩИ 5). o5 5050500008 Toil 18.1 85.4 29.2
William's Чезроай. ое... Me ies а... 5.5 9.5 | 2.0 | 20.8
TNEMBCAMNCE SIAM mole WO 55555065 n000050 5,0 Absent
Milsraronysalmasremr ry Не, о ee clei 4,4 42.4 2.9 12.5
Sima tive=toecdigerbod) |. р. ene A) 0.2 0.3 0.3
European Naremepeg. ; осо он 2.2 = 0.6 Зо
Азта MinoOmiamisten <0 0) © . се. 1.6 Absent
Comin weil ИО о о ов оо ово bo! 1.6 } 5 | 0.2 | =
ЗЕРНО Е ее иены: ne То Г 3.2 —
Пао ИО лы р ао о ооо О 1.1 Absent
Рог 60199758 ооо бло сю Gnd oun He воно 0.5 = 0.1 =
INKOLAWENY ENE ооо о або о Galo © oda myo Bila orc = = 0.1 ==
ПМТ NS OF GXSCIES 55 ор обо co blo dn eo 8 15 7 9 6
Number ое ха а О tees sels 181 347 2,473 24
* Recent Indian porcupine is а possible descendant of the Binagady species.
TABLE 24. Changes in composition of Quaternary Perissodactyla in the piedmonts of the eastern Caucasus
Pleistocene
(150)
Historical time
Species
Ree Apsheron and Dagestan
Apsheron ; ;
Kabristan piedmont
FEM WS IGP UIMEN MUS > o6ca0. von accor =
2, MCMV OMUS eon o do 6 ROHS Lee cet eee =
в, сааб yo 5 55560 a0 on ar
Е, CAAIMIS SUDA, 55385000000 0
RIMUNIGPC EROS lylme saGdeMEnS 55546.
152 In studying the Pleistocene landscapes of the Apsheron, birds contribute
little to our knowledge because of their seasonal migrations. Beetles,
however, are more reliable. The proportions of beetles found at Binagady
in aquatic, amphibious and terrestrial habitats respectively is given in
Table 26 (identification by A.V. Bogachev; author's figures).
Aquatic beetles, as such, are not indicative of the landscape, because
of their capability of flying great distances; water-scavenger beetles and
water beetles have even been known to cross great expanses of desert to
reach water. According to Bogachev, land beetles in the Binagady
bone -bearing bed comprise 8% (out of a total of 75.7%) of relatively
mesophilous forms which are characteristic of the present piedmont brush
near Shemakha and are absent from contemporary Apsheron. In other
145
(151)
FIGURE 70, Binagady
1-4—M?, М! heel bone and hoof of Equus caballus subsp,; 5-9 —М2, м, Ртз-М» heel
bone and pelvis of Equus hidruntinus; 10, 11 —skull of Rhinoceros binagadensis
(top and bottom views)
146
(150) TABLE 25. Changes in composition of Quaternary Artiodactyla in the piedmonts of the eastern Caucasus
153
Historical time
Apsheron
Е
pecies Apsheron and Dagestan
Kabristan piedmont
SOG LSC eeoNy OWS he aio odes ooo WO
а.о. o.oo ogee
Cervus elaphus
Megaceros euryceros
SIGE Бата Ge ooode pce nooo Oe
Gazella subgutturosa
Ovis cf. ammon
Bos mastan- zadei
Be ри оао е п о. cs oe Reem 2 ?
оон
Bison bonasus
3х
It is possible that the Apsheron boar is ancestral to the contemporary species,
** The primitive bull inhabited Apsheron later than the period of the Binagady trap.
words, the shift in the range areas of beetles confirms a progressive
development of xerophilous landscapes in the Holocene.
Thus, both qualitative and quantitative data on the mammal and beetle
fauna and on the flora show that the climate of the Pleistocene Apsheron
was colder and moister than in the present. The ecological conditions
were Similar tothose which now exist in the thinly forested, arid zone
on the southern slopes of the Armenian Highland at elevations of 1,500-
1,600 m, and in the Dagestan piedmont at elevations of 0-500 m.
The universality of the Binagady assemblage leads to other conclusions
regarding the stratigraphy, paleoecology and history of the formation of
the fauna.
The Binagady faunal complex contains a representative group of species
widely distributed in the Pleistocene over steppes and forest-steppes, and
known from Paleolithic and other localities of Europe. These include:
Erinaceus europaeus Apodemus sylvaticus
Canis lupus Mus musculus
Vulpes corsac Microtus arvalis
V. vulpes Arvicola terrestris
Crocuta spelaea Equus caballus
Ursus arctos Megaceros euryceros
Meles meles Cervus elaphus
Panthera spelaea Saiga tatarica
Allactaga jaculus
The origin of most of these species is not yet clear. Oniy the corsac fox,
great jerboa and saiga migrated from the north into eastern Transcaucasia.
From the ecological point of view, the above assemblage is heterogeneous.
It includes some species which, at present, are adapted to open habitats,
and some species which live in the forests and moist river valleys of the
same regions. There are no true xerophilous forms in the assemblage.
Some of the listed species (fox, wolf, bear, badger, great jer>doa) are
represented at Binagady by southern forms, an index of the antiquity of
the conditions under which the Binagady faunal complex evolved.
147
(152)
FIGURE 71. Binagady
1,2 — epistropheus and metatarsal of Sus apscheronicus; 3,4—metatarsal and humerus
Ovis aff. ammon: 5-8—horn, atlas, scapula and metatarsal of Saiga tatarica binagadensis;
9, 10 —skull of Bos mastan-zadei, from above and side (1/20 natural size) (from Bogachev, 1940)
TABLE 26, Ecological groups of beetle species and numbers of individuals from the Binagady asphalt
Number of individuals
Number of species
15
11
81
Ecological groups of beetles
© ом MMe) elie 6 оо о за ею
The species which became extinct are: cave hyena, cave lion,
horse, giant deer. The ranges of corsac fox, great jerboa and saiga
have shifted considerably, but only localized changes took place
in the distribution areas of the common and water voles.
154 There is a prominent group of Mediterranean species adapted to
xeromorphic landscapes, such as the upland steppes of southwest Asia.
These species are:
148
155
Crocidura russula Microtus socialis
Hemiechinus auritus Ellobius lutescens
Vormela peregusna Meriones erythrourus
WEIS lytic Hystrix vinogradovi
Acinonyx jubatus Rhinoceros binagadensis
Lepus europaeus Equus hidruntinus
Allactaga williamsi Sus apscheronicus
Cricetulus migratorius Ovis ammon
Mesocricetus auratus ‘ Bos mastan-zadei
The group includes species adapted to mixed biotopes (white -toothed
shrew, long-eared hedgehog, tiger polecat, jungle cat, cheetah,
William's jerboa, steppe vole and boar), to upland steppes (Radde's
hamster, migratory hamster and Transcaucasian mole vole), and to hot
valleys and lowlands (red-tailed gerbil and Binagady porcupine).
Most of these Mediterranean forms were represented in the Pleistocene
by distinct subspecies, whose main areas of distribution at that time lay
definitely to the south of the Caucasian Isthmus. The extinct Pleistocene
ass (Equus hidruntinus), Binagady rhinoceros and Binagady goat were
also of southern and southeastern origins. The extinct Apsheron vole,
which has been identified from its molar classification as belonging to the
European type of subgenus Pitymys, can be included among the endemic
species. The range of the Dagestan hamster shifted into inner Dagestan,
and that of the mole vole, to the mountain-steppe of Talysh.
Thus, the general evolutionary trend of the faunal complex on the eastern
Transcaucasian plains was toward either extinction or shifts in distribution
patterns of relatively mesophilous forms. The disappearance of these forms
from the area was caused by the development of a drier climate and
consequently of a more xerophilous landscape.
The general paleozoogeographical relationships within the Binagady
complex can be understood when the complex is compared with the
Pleistocene faunas of the southern part of the Russian Plain and of Southwest
Asia.
Some common species of the Middle and Upper Pleistocene of Eastern
Europe do not occur at Binagady (e.g., mammoth, woolly rhinoceros,
Bison priscus, cave and other bears). Northern species were also
absent from the Binagady complex: arctic fox, blue hare, lemming,
reindeer, musk-ox.
The absence of these forms may be accounted for in part by the existence
of a xeromorphic landscape in the eastern Caucasus, and in part by the
geological age of the locality.
Tundra species probably did not extend as far south as the area of the
Binagady trap, at the time it existed. Neither the harvest mouse, striped
field mouse nor Siberian polecat reached the area from the north. And so
it was that only some of the faunal elements of the southern Russian Plain
and of the Iranian-Asia Minor uplands met in Apsheron and in Kabristan
during the Pleistocene. Certain southwest Asian species, listed below, which
have been found in the Middle and Upper Pleistocene of Palestine, Syria
and Lebanon, were also absent from the area under consideration:
149
156
Herpestes ichneumon L, Dama mesopotamica L,
Felis chaus Guld, Gazella arabica L.
Panthera pardus Schreb, Gazella cf, subgutturosa Сша.
Rattus rattus L, Capra primigenia Fraas
Spalax sp. 7 С. beden Schreb,
Phacochoerus garroda Bate Bubalus sp,
Hippopotamus amphibius №. Procavia cf, syriaca Schreb,
The distance between Apsheron and Syria and Palestine indicatcs that
the differences in composition of the fauna have zoogeographical significance
which persisted through geologic time.
A comparison of the Binagady complex with the Recent fauna of the eastern
Transcaucasian plains shows the absence in the earlier period of a number
of eurytopic species — species which probably penetrated the Apsheron at a
later time, some still extant, suchas Suncus etruscus, Canis
aureus, Hyaena hyaena, Mustela nivalis, Felis chaus,
F. lynx, Panthera pardus, Rattus norvegicus, Gazella
subgutturosa and Martes foina, and some recently extinct, such as
Panthera leo and Equus hemionus.
Thus an interesting feature becomes apparent: a number of southern
thermophilous species migrated to eastern Transcaucasia, while a number
of European steppe species retreated into eastern Ciscaucasia in the period
following the time of the Binagady trap.
All of which emphasizes the specific features of the Binagady complex
and the relatively short period of time during which the asphalt trap was
operative in the Pleistocene.
Stratigraphic correlations and age of the Binagady complex
The age of the Binagady locality is significant in tracing the origin of
Quaternary fauna and the morphogenesis of mammals.
Attempts at dating the bone-bearing bed and the faunal complex of
Binagady have been based on the estimated age of the old Caspian terrace
and of the orogenic compression of beds which was accompanied by an oil
flow, and on paleozoogeographic correlations.
Assuming the age of the terrace underlying the bituminous beds to be
Khazar, Bogachev (1939, pp.47-51) has dated the Binagady locality Riss-
Wurm. He has also related the compression of the procuctive beds and the
oil flow to the so-called ''Kalinovka"' phase of folding. According to Reingard
(1937) this phase occurred at the beginning of the second Riss glaciation.
According to Vardanyants (1948, table 3, pp.104, 125), considerable
compression occured in the pre-Khazar and pre-Khvalynsk orogenic phases,
and later in the Neo-Pleistocene and Holocene, resulting in the formation
of folds on the Apsheron and the development of mud volcanos. According
to this scheme, the Binagady bed might have formed either in pre-Khazar
or pre-Khvalynsk time. , .
Bogachev places the tirne of truncation of the Binagady fold at the Khazar
transgression, and concludes from this that the oil which appeared during
the Kalinovka orogenic phase could not have been instrumental in trapping
and preserving plants and animals, because the oil would have disappeared
150
157
prior to the Khazar transgression. Accordingly, Bogachev (1940Ъ, р. 14)
postulates the dislocation of Didacna surachanica beds and the
existence of two Kalinovka compressive phases.
An assumption that the Binagady fold was truncated by the Baku sea
would, in fact, lead to a correlation between the formation of the asphalt
trap and the pre-Riss and the Kalinovka orogenic phases, and would affect
an estimate of the age of the locality.
The peculiar features of the Binagady complex, as compared with the
Quaternary complexes of the Russian Plain, complicate biostratigraphic
correlations. Moreover, the difficulties are compounded by the lack of
index fossils, such as elephant and bison, at Binagady.
Gromov (1948, p.424) has dated the Binagady complex Upper pre-Glacial,
correlating it with the Middle Quaternary ''Khazar fauna of the Volga
region,'' which Nikolaev (1937) dated Mindel-Riss.
The Burchak-Abramovich —Dzhafarov summary (1955) indicates that a
majority of paleontologists accepted the Bogachev estimates without verifying
them independently.
In a study of carnivores from the Binagady asphalt (Vereshchagin, 1951b),
the author employed the same age estimate with a disclaimer that the dating
can be improved through more complete excavations and further study of
the relationships of the continental and marine beds at the Binagady mound.
A comparison of the Binagady species with lists of species from the
Paleolithic caves of Palestine, Syria and Lebanon (Bate, 1937; Picard,
1937) shows that the greatest similarity between the faunas of the two regions
existed during the Mousterian and Aurignacian, i.e., in the Upper
Pleistocene.
We determined the residual organic content of the Binagady horse and
saiga bones by calcination (as used by Pidoplichko, 1952). The results
showed that upon oil extraction by hot alkali and benzine, the calcination
index was in the 480-520 range, which is close to the index of the bones
of the Khazar complex.
V.V.Cherdyntsev's attempt at a radioisotope determination of the age
of the bones was unsuccessful.
On the basis of all the paleobiological and geological data, the Binagady
complex must have existed and been buried at the beginning of the Khazar
transgression, i.e., the end of the Middle Pleistocene. In Reingard's
scheme, this corresponds to the lower part of the Middle Caspian beds and
the second Riss glaciation (''Ry''); in Gromov's, it corresponds to the
Riss, "QL".
How long the Binagady complex existed within the framework of the faunal
composition described is not accurately known. Some species of the complex
continued to live even in the Khvalynsk age, while others were replaced.
An example of a later deposit where the Bos of the Asian type (Bos
mastan-zadei) shows replacement by the European tur, while the saiga
continued to exist, is another asphaltic locality, Kir-Maku.
Morphological development of eastern Transcaucasian animals during
the Upper Quaternary is discussed in Chapter IV.
The results of the taphonomical and zoogeographical analysis of the
Binagady burial may be summarized as follows.
1. The Binagady mammals lived in a dry steppe and in sparse, arid
forests which grew in gulleys and in areas of placer rock. The climate of
the region was somewhat cooler and more humid than in the present.
151
158
2. Animals of various ecological types perished in the liquid oil and
viscous asphalt, as well as in mudflows.
3. The animals were buried in the asphalt sand and fine-grained coquina
at or near the site of death. The asphalt beds were later covered by eolian
sandy coquina and lacustrine-estuarine sediments.
4. On the basis of the occurrence of Khazar molluscan index-species
in the Early Caspian terrace, the Binagady bone-bearing bed on the terrace
margin can be dated post-Baku age.
By relating the profuse oil flow and the deposit of animal remains with
the orogenic phases, the Binagady faunal complex can be dated as either
Lower Khazar or Lower Khvalynsk.
5. A comparison of the Binagady faunal complex with the Quaternary
complexes of the xerothermal zone of southwest Asia (the Paleolithic of
Syria, Lebanon, Palestine) indicates that the Binagady complex correlates
with the Mousterian culture. The coefficients of the residual organic matter
of the Binagady bones are similar to those of the Khazar bones of the Volga
region.
6. Of the 39 mammal species in the Binagady complex, 19 (48.7%) are
Pleistocene species found widely distributed on the plains of Western and
Eastern Europe. Another 19 (48.7%) are Pleistocene species of southwest
Asian and Iranian origin. Only the one remaining species (2.6 %) can
possibly be endemic.
The faunalcomplex probably did not include arctic or mountain animals,
or cave bears, elephants andbison. Duringthe time of the complex, the
ranges of several animal species of the steppes of the Russian Plain and
of the upland steppes of southwest Asia met on the Apsheron. Fossiliferous
localities of younger age on the Apsheron contain faunal assemblages which
are not as universal as that of Binagady.
Kir-Maku
The locality is 12 km south-southeast of the village of Dygya, in
the center of the Apsheron Peninsula, on the slope of the Kir-Maku asphalt
mound.
The Kir-Maku site was discovered in 1870 by Shtukenberg, who
collected and gave to the Kazan University a skull of a female saiga anda
wolf mandible in an asphalt matrix.
According to the map of Volarovich and Lednev (1913-1929), the site
lies 100 m above sea level; the mound is nearly 1.5 km wide at the base
and is 60-70 m high. The top of the hill is composed of Early Caspian
deposits, underlain by almost vertical middle productive sands (Middle
Pliocene). Two buried or half-buried asphalt flows lie on the northern slope
of the hill; one asphalt flow occurs on the southeastern slope. A bone-
bearing kir layer occurs somewhat below the top of the hill, at a depth of
2-5 m and covers an area of 1.5-2 hectares. A generalized stratigraphic
section through the bituminous beds of the Quaternary at Kir-Maku as given
by Sultanov (1947) is shown in Figure 72.
$52
159
The bitumen in this area often occurs in vertical veins, nearly 20 cm
thick at the base and 1.5-2 m long. However, the main mass of the bitumen
lies on a continuous layer, probably of flow origin. Bones occur in both
the stratum and the veins. The oil which was forced to the surface by
compression of the beds probably formed sticky crusts and pools which were
later covered by sand and dust. Only fragmentary skeletons and skulls are
found, as arule, inthe bitumen, probably remains of animals that died
trapped in the asphalt crusts.
The Kir-Maku locality has something in common with the
travertines of Mount Mashuk in the northern Caucasus. However, the
asphalt crusts probably acted as a more efficient trap.
Bogachev (1924, 1925b) reported a skull
of Bos primigenius that was
iam 8 7972 ол , indistinguishable from a species which lived
а Е оп the Russian Plain during the Pleistocene
and Holocene. He dated the bituminous salina
near the village of Dygya post-Baku and
thought that the Bos died and was preserved
during a period of increasing humidity and
expansion of the steppe flora on the Apsheron
Peninsula in Khvalynsk time.
In 1940-1941 a large number of bones of
Bos primigenius, wolf, fox, cave
hyena and vulture were found in the process
of bitumen-mining. The miners buried the
bones in a specially prepared trench
(Bogachev, 1944; Sultanov, 1947).
i i). The Kir-Maku beds are younger than the
Binagady beds. The age of the relief at
Kir-Maku can be readily determined, since
the bituminous bed is concordant with the
slope.
Thies BOS реет oe mss) оао
Maku is a younger type which replaced the
Binagady form in the Upper Pleistocene of
eastern Transcaucasia. The Kir-Maku
Pleistocene assemblage still included saiga
and possibly hyena of the Binagady type.
Fossils found in the bitumens of the
northeastern part of Artem Island belong to
the Upper Pleistocene. These include two
red deer skulls with broken antlers which
FIGURE 72. Section through the asphat- | Were placed in the Museum of Natural
bearing beds at Kir-Maku History in Baku (Bogachev, 1938a). Other
ae. es Си HOS saneyMeliyoy bones of red deer, and several bones ofhorse,
sequence; с — “rich” asphalt bed; d — wolf and large birds were collected at the
productive sands, On the right — same locality in 1944 from old caches
thickness in meters (Burchak-Abramovich, 1951с).
The age of the bones from the island
approaches those of Kir-Maku, as nearly
as can be inferred from the state of preservation of the material.
SS)
HISTORICAL EPOCH Greater
Caucasus
Broadleaf forests
Saline semidesert Alpine zone
of uplands Piedmont
semisteppe
Ephemeral Shemale Mount Baba-Dag
semidesert a
Binagady 3000
< | ; we 2000
| Ся.
0
MIDDLE PLEISTOCENE
Greater
Caucasus
Broadleaf forests
ee Piedmont Alpine zone
Binagady locality pela Везер Be 3000
р 2000
еже —t
ia 0 10 20 30 40 50 60 70 80 90 100 N10 120 130 140 150 160 km
FIGURE 78. Changes and migration of landscape zones since the Middle Pleistocene on the Apsheron
Peninsula, based on the study of the Binagady locality
Deer and horses probably lived on Artem Island during the relatively
humid, cold period inthe Top Pleistocene. At that time ungulates migrated
freely over the Apsheron and to nearbyislands, particularly during the
spring growth of ephemeretum. Occurrences of animals on the islands,
therefore, do not indicate a regression of the Caspian, since the animals
could swim across the narrow straits.
The Pleistocene development of landscapes and faunal complexes on the
eastern Transcaucasian plain can be described from data at hand, as
follows.
The development of the xeromorphic landscapes which predominated
in the Lower Pleistocene probably began in the Middle Pliocene (the time
of the productive beds). During the Baku transgression the shores of Kura
bay were inhabited by rhinoceroses, Elasmotherium, horses, camels,
and, of the carnivores, hyenas which closely resembled the spotted or
African brown hyena.
160 A distinct complex of mammals developed on the plains of eastern
Transcaucasia toward the Middle Pleistocene, consisting of species widely
distributed on the steppes of the temperate zone of Europe and on the
upland steppes of southwest Asia.
154
161
By analogy with western Transcaucasia, it can be assumed that the
mountains of the Bolshoi Range were inhabited by European brown bears
and cave bears, martens, lynx andherds of mountain goats and chamois.
Glaciation over the eastern spurs of the Bolshoi Range, while probably
not extensive, was sufficient to create a number of changes in the fauna
of the plains. The general landscape was characterized by sparse
forests and steppes.
The Bos of the Asian-Indian type was replaced by the European tur.
Argali and giant deer probably disappeared. A number of Pleistocene
species of the steppes and forest-steppes of Eastern Europe penetrated
Transcaucasia, and possibly the Iranian Plateau, during a cool, humid
climatic phase in the Upper Pleistocene. During this stage the altitudinal
vegetation zones probably migrated down the mountain slopes (Figure 73).
This, however, did not interrupt the continuity of the xerophilous fauna
of southwest Asian origin.
The East European steppe animals (corsac fox, great jerboa, saiga)
retreated to the north only at the very end of the Pleistocene during a new
xeric climatic stage.
A new migration of southern species into Transcaucasia occurred at
the beginning of the xeric Holocene epoch. Some of the species migrated
as far north as the Ciscaucasian plains.
Lesser Caucasus upland (southern
Transcaucasia)
The high plateaus of southern Georgia, Armenia and Karabakh were
taphonomically in a special position in the Pleistocene. Frequent mudflows,
landslides, rockfalls and volcanic eruptions, associated with the
mountainous terrain, caused death among animals. However, their incidence
as fossils is rare. The known Pleistocene localities are situated on the
margins of broad river valleys and in sink holes of high plateau lakes. The
bones were deposited in river and mudflow fanglomerates, in diluvial
sediments near mountain slopes and in lacustrine sediments.
Although Paleolithic material is plentiful on the Armenian Highland,
the instrumentality of man in the accumulation of animal bones was
negligible (Zamyatnin, 1947; Panichkina, 1948, 1950a, 1950b; Sardaryan,
1954), probably because of the erosion of most of the Middle Paleolithic
caves and open sites.
A site containing bones of Lower Pleistocene mammals in ancient
lacustrine deposits near Akhalkalaki is particularly interesting. The
locality was discovered by Vekua in 1958 and from it he has identified the
following species: Canis sp., Crocuta spelaea, Meles cf. meles,
Marmota sp. (smallform), Elephas cf. trogontherii, Equus
sp., Rhinoceros etruscus, Hippopotamus sp. (large form),
Megaceros sp., Bos cf. primigenius. The presence of the
hippopotamus and of an unusual marmot in the assemblage emphasizes its
southern aspects. The incidence of both forms indicates that the landscape
of the area at the dawn of the Anthropogene was comprised of low plateaus
with a well-developed river network and warm climate. *
* The age of this assemblage may be older, possibly Upper Pliocene.
155
FIGURE 74, Skull of Bos primigenius from Bayandur on Armenian Highland
A large accumulation of bones was discovered in 1928 in the sand-gravel
quarries near the Kazachii post on the southern outskirts of Leninakan, of
which up to 300 kg were removed to the Yerevan Museum (Bogachev, 1938c,
1938d; Avakyan, 1948). Bogachev has identified the following species in
the material collected: Bison priscus, Megaceros sp. (hiberni-
cus?), Cervus elephas maral, Equus caballus. Bones,of an
elephant (Elephas cf. armeniacus) were taken from a lower bed
which has been assigned to the Upper Pliocene (Apsheron age or equivalent).
The rest of the bones were dated Pleistocene. Avakyan and Burchak-
Abramovich (1945) have identified the camel Camelus knoblochi in
the material and the following species occurring with it: Equus
stenonis, Rhinoceros cf. mercki, Elephas meridionalis,
E. trogontherii (?), Cervus sp., Bos sp.
Elephant teeth from near Leninakan, described by Sardaryan (1954),
indicate that the predominant species in the collectionis Elephas
trogontherii, which closely resembles the Tiraspol E. wusti and,
to a lesser extent, the elephants from the Taman Peninsula conglomerates.
A fragment of skull of Bos primigenius from this locality (collections
of the Museum of Georgia) has been described by Burchak-Abramovich
(1951d).
Another fragment of Bos primigenius skull (Figure 74), taken from
near the village of Bayandur in the Leninakan area, has been described
by Gromova (1931). Even this brief review of the material clearly shows that
fossils in the alluvial deposit near Leninakan are of varying ages. The
deposit strongly resembles the gravels of the inclined Ciscaucasian plains.
A Lower Quaternary faunal complex, with such indicative forms as
Elephas trogontherii, Equus stenonis and Rhinoceros
mercki, is distinguishable in the material collected and shows acorrelation
of the Leninakan freshwater deposit with the upper part of the Psekups
gravels and the lower beds of the Girei quarry on the middle Kuban.
The Lower and Middle Quaternary bones from the sands and gravels
near the village of Eilas, 12 km southwest of Yerevan, are close in age to
the material described. In the Adzhi-Eilas site the gray, fine-grained river
sands containing mammalian bones underlie the 3—3.5-m-thick gravels and
loessial loams. The bone-bearing lenses are up to 1-1.5 m thick. Some of
the bones had been redeposited, being apparently derived from older
lacustrine sediments, The beds were dated as Riss age by Avakyan (1946).
The fragments have lost most of their organic matter. The surrounding
matrix of the horn fragments is ''chalky'' and ''marly.'' The enamel of the
deer teeth is grayish brown from staining of the dentine. The material which
1704 156
162 We studied includes teeth of Elephas trogonthetii, a fragment of
the skull of ''diluvial-type'' Bos, Bos trochoceros, fragments of deer
horns and teeth, Cervus ex. gr. elaphus and Dama cf. mesopota-
mica (Figure 75).
FIGURE 75, Adzhi-Eilas
1 — skull of Bos trochoceros; 2, 3 — fragment of horn and jaw of Dama cf,
mesopotamica
Younger, Upper Pleistocene bone deposits, partly created by primitive
hunters, are so far known only in the northern sections of the Lesser
Caucasus upland. There is a record of rodent bones, a fragment of a deer
horn, fragments of tusks and skeletal bones of Elephas primigenius
and a tooth of Equus caballus taken from a depth of 10 m at a site
on the left bank of the Chichkhana rivulet near Nalband, 30 km north of
Leninakan (Karakash, 1898).
Zurtaketi site. This Paleolithic site discovered by Maruashvili
(1946) on the Tsalka Plateau in southern Georgia was ascribed particular
stratigraphic significance. The site is located at an elevation of nearly
1,320 m on the right slope of the Karabulakh River valley (the right
tributary of the Khram River), 15 km north of Bashkicheti. Fractured bones
and tools of flint, obsidian and bone reflect human activity at the site. The
obsidian tools are ten times as abundant as the flint implements. The
bone and stone tools must have been attached to handles, because they
157
163
resemble darts and arrowheads. From the stone material the site has been
dated as Solutrean,
Because he assumed that the primitive tribe was driven off the site by
a lava flow which burned and covered the cultural layer, Maruashvili thought
that the uplands relief and the volcanic activity dated from a period later
than that of the site. His age estimates were disproved by Paffengol'ts
(1951) who established that the site is located in a collapsed recess ofa
doleritic lava wall.
The mammalian material collected by Maruashvili in 1945 and 1948
consisted of molars, incisors and metatarsal epiphysis of a horse, molars
of an ass, vertebrae of a proto-bison and a fragment of a ram radius.
Most of the bones are weathered and have slightly dendritic surfaces.
Their state of preservation indicates a very late stage of the Pleistocene.
The material from the Zurtateki site, as identified by Burchak-
Abramovich (1951а), is given in Table 27. In addition, partridge bones
which resemble rock partridge have also been identified in the locality
material.
The species composition of the game of the Zurtaketi people reflects a
Khram Basin landscape of rolling plateaus covered with meadow-steppe,
TABLE 27, Mammalian species and number of bones from Zurtaketi site
Species Number of bones | Number of individuals
Equus caballus subsp. ......
Е, (Asi muss:) Sp. ЗВ be tote ses
Bia SOM PLS DUS! а ее
Onis, Свое iM. weyers eae
RRR о
ese sie пая мае
and underdeveloped forest-steppe vegetation in the Upper Pleistocene.
The asses (kulans) and small rams were quite common until recently
in the woodless territories farther south.
Prehistoric hunters were also active along the upper Khram River,
where a Mesolithic cave site, approximately 8 m above the riverbed, was
discovered by Kuftin (1941) near the village of Barmaksyz.
At this site, a cultural layer of no more than 40 cm in depth yielded
9,900 broken obsidian tools and 600 flint tools in a 70 гп? area of eroded
slope with surface soil up to 80 cm thick.
The bone material consists of fragments of diaphyses and ungulate teeth,
stained with humus.
According to Burchak-Abramovich (1951b), the collections include six
molars of a small horse and a lower molar of a Bos.
During the artificial lowering of Lake Sevan, Dal' (1950b) collected
material along the shores which is the same geologic age as the collections
mentioned above.
The extensive collections are mostly fragments of bones of domestic
animals of post-Paleolithic age which are discussed in more detail below.
As far as the state of preservation shows, the oldest bones belong to a
Bos cf. minutus and adromedary (Camelus dromedarius),
164 tentatively identified by Квауезоп (1954b) as a wild form.
158
165
The fauna of the Pleistocene beds of southern Transcaucasia is relatively
poor in species. Nevertheless, there is a regularity in the replacement of
the faunal complexes one by another, similar to the process which operated
in Ciscaucasia.
It is interesting to note that a number of index species characteristic of
the ''cold steppes" of Europe occur in the Pleistocene deposits of the Lesser
Caucasus plateau. The following species are known: Elephas trogon-
ое об VO Cie Oise Ime inc kul ое ате 5 као ось,
Megaceros, Cervus elaphus, Bos primigenius. Of the species
peculiar to southwest Asia, only fallow deer and mouflon-like sheep are
known. The southern east Mediterranean aspects of this faunal complex
were probably better developed among the smaller forms in the Pleistocene.
The species under study have not yet been classified stratigraphically,
nor have morphological characteristics been established for the index forms
which take into account their environmental adaptations.
As opposed to the Greater Caucasus, the Upper Pleistocene faunal
complex of the highland region under discussion has not yet been recorded.
This is partly accounted for by the poor development of the karst, and by
the fact that the karst itself has been inadequately studied. In the face of the
scarcity of Paleolithic material, it is impossible to form a judgment of the
effects of Pleistocene cooling and glaciation of the plateaus on the
development of the fauna. There is no doubt, however, that the increase in
water and in mesophilous vegetation on the plateau did not preclude the
existence of camels in the hot intermontane valleys. The vertical shifts of
vegetation zones and faunal complexes which occurred during the period of
cooling in the Pleistocene were probably less extensive than in the Greater
Caucasus.
The period of transition to the Holocene on the Armenian Highland is not
very well known; it is probable, however, that it was similar to that in
eastern Transcaucasia.
The stratigraphic and geographic distribution of Pleistocene mammals
in the Caucasus is given in Figure 2 (see Introduction).
ECOLOGY AND LANDSCAPES OF THE CAUCASIAN
ISTHMUS ACCORDING TO PALEONTOLOGICAL DATA
A review of the geological data and of Pleistocene fossiliferous localities
by regions provides a basis for a summary of the specific features of the
development of Caucasian fauna.
From the excavations at Kudaro I, the faunal complex which existed in the
mountains that were uplifted in the Lower Pleistocene can be recognized
as the direct ancestors of Recent Caucasian species of insectivores, rodents
and ungulates which are characteristic of the Mediterranean region of the
Alpine folded belt. From the zoological point of view, the strong
peneplanation of the Caucasus in the Lower Pleistocene which is assumed
by some geologists (see, for example, Vardanyants, 1948)is an
impossibility. The evolution of the Caucasian mountain fauna with su¢éh
highly specialized forms as Prometheomys, Microtus nivalis,
Caucasian goat and chamois took place at the same time as the evolution of
similar forms in the Alps, the Carpathians and Asia Minor. It is difficult
159
166
to support a case for the origin and development of these mountain species
only in the Pleistocene, since, by the end of the Middle Pleistocene, they
had already assumed characteristics which are identified with the Recent.
The main sections of the Greater Caucasus which had been uplifted during
the Miocene probably retained their elevation into the Pleistocene, even
though they might have been somewhat eroded.
The better-known death assemblages of Middle and Upper Pleistocene
mammals in the Caucasus reflect a continuing cooling of climate and
modernization of fauna.
Elasmotherium and Cervus pliotarandoides disappeared
in Ciscaucasia and Transcaucasia toward the Middle Pleistocene. Steppes
and forest-steppes replaced the semideserts and savannahs on the piedmont
plains and the evolution of landforms promoted further isolation of faunal
complexes in different landscape zones. At that time, the forest-steppes
of Ciscaucasia were inhabited by Elephas trogontherii, Rhino-
ceros mercki, red deer and long-horned bison. In eastern and southern
Transcaucasia the bison were replaced by Bos primigenius.
A widespread invasion of eastern Ciscaucasia by some xerophilous
species of the Aral-Caspian fauna (e.g., little suslik, Central Asian gerbil
and jerboa) might have occurred very early, perhaps even at the close of
the Apsheron. It is possible that the ancestors of these species survived
the Caspian transgressions on lands bordering the east Ciscaucasian bay,
or that, having disappeared from the area, they returned in new migrations
during the regressions of the sea.
A mountain-forest faunal complex, consisting of bear, boar, deer,
Caucasian goat and bison, existed in western Transcaucasia on the steep,
southern, seaward slope of the Bolshoi Range.
The faunal migrants from the Russian Plain were represented by the
common hamster which appeared as far south as Tsebel'da.
The low plateaus of Imeretia formed the northwestern boundary of the
distribution zone of Upper Pleistocene xerophilous species of the southwest
Asian uplands: Radde's hamster, porcupine, ass, argali-like sheep. Apes
also survived in the area from the time of the Pliocene. The presence of
horses and asses indicates a steppe development along the middle section
of the Rion.
Similar steppes, covered in places by very extensive lava flows, also
developed in the Lesser Caucasus uplands and in central Transcaucasia.
At the end of the Lower Pleistocene, the large mammalian fauna first
showed noticeable effects of Paleolithic man's presence in the environment.
These effects were more pronounced in the western part of the Isthmus.
Commencing in Upper Pliocene time, the eastern Transcaucasian
landscape, composed of dry foothills, juniper-pistachio forests and steppe
grasses dry in summer, persisted throughout the Middle Pleistocene. The
faunal complex of this region had a mixed composition of animals from the
Pleistocene steppes of the Russian Plain (corsac fox, saiga, etc.), from
the southwest Asian uplands (jungle cat, cheetah, mole vole, etc.),
and from deserts of the Turan type (small five -toed jerboa). It also included
endemic Caucasian species (Apsheron vole).
As a general rule, cave bear, elephant and bison were not to be found.
The only northern steppe species from the Pleistocene Russian Plain to
occur in eastern Transcaucasia during the Middle Pleistocene were corsac
fox, great jerboa and saiga (Figure 76).
160
= INFERRED AREAS OF
asp DIST RIBUTION OF
MAMMALIAN
END OF THE MIDDLE
PLEISTOCENE ON THE
q CAUCASIAN ISTHMUS
(72
Us __ 7
US ty
777 12:
ный = To
! hl Se №
_
Ht ali
=\
——
——— 55
р Caucasian mountain- South Russian steppe
727 [aa]
forest and mountain- complex
meadow complex ЕР) Central Asian semidesert
(mo Southwest Asian complex
mountain-steppe complex
FIGURE 76
The ranges of the "northern" and ''southern' steppe species expanded
along the margins of the Isthmus toward each other, probably moving closer
together in irregular waves —irregular, that is, in the sense of both space
and time. It seems possible that the extension over the Caucasus of the
ranges of some other northern steppe species (e.g., common vole, Bos
primigenius) also proceeded unevenly even in the Middle Pleistocene.
The process was controlled by the alternation of arid and humid climatic
phases.
The proportion of faunal elements from the Russian Plain steadily grew
throughout the Pleistocene. The effects of this incursion of Russian Plain
species was more pronounced in the fauna of the Ciscaucasian plains than
in that of the Transcaucasian.
True mammoths and bison, but of somewhat diminished size, lived in the
western Ciscaucasian foothills at the beginning of the Upper Pleistocene,
at a later time than that of the Binagady complex.
161
167 The extensive development of steppes оп the Trans-Kuban Plain at the
beginning of the Upper Pleistocene can be seen in the occurrences of grass
beetle, horse, ass and saiga remains at the Il'skaya site. Mammals from
the Upper Pleistocene localities of the Ciscaucasian plains also indicate
the degree of development of the steppes. The mammals of this region
include horse, boar, red deer, saiga, bison and Bos primigenius.
Such an assemblage may be characteristic of either a steppe or a forest-
steppe, but not of extensive forests nor of mesophilous meadows, and
certainly not of moss swamps and tundra.
From available paleofaunal data, it appears that glaciations in Eastern
Europe and on the Caucasian mountains did not noticeably affect the
composition of the faunal complexes on the piedmont plains. Moreover,
development of desert and semidesert landscapes, rather than mesophytic
landscapes, is indicated by the large collections of bones from the shoals
of the lower Don. The occurrence of Upper Pleistocene and Holocene bones
of blue hare and reindeer along the lower Don must be regarded as
evidence of a late extension of their ranges southward along the valleys of
the larger rivers of the Russian Plain, and not as a direct shift in the
distribution areas caused by the cooling and glaciation of northern Europe.
Mountain forms, such as snow vole, Caucasian goat and chamois, are
completely unknown in the Pleistocene of the Caucasian foothills and plains.
This might be taken as a contrary indication to the assumptions of
continental glaciations and major shifts in the phytolandscape (such as
displacement of alpine groupings to the plains) held by earlier zoologists
and contemporary geomorphologists.
If, during the glaciations of the Caucasian mountains, the glaciers
descended to the piedmont plains, then the highland fauna (mountain goat,
chamois, Prometheomys) must have either been completely displaced
to the piedmont plains, or migrated to some adjacent mountain refuge,
or become extinct.
The cooling and the concomitant down-slope movement of the altitudinal
zones can be traced through paleontological material which reflects the
wider distribution of mesophilous animals and animals of the mountain-
forest zone from the time of the Middle Pleistocene in western and eastern
Transcaucasia.
It should be noted that the Caucasian Isthmus, because of its more
southerly location and more varied relief, had a more variegated species
population during the Pleistocene than the southern part of the Russian Plain.
Evidently, during the Pleistocene a number of subtropical and hydrophilous
species of mammals found refuge in Transcaucasia. A number of such
forms, such as macaca, black rat and porcupine, survived through the
Pleistocene in Colchis and Asterabad because of proximity to the Black Sea
and Caspian basins.
All of this is fairly well in agreement with the results of paleontological
studies in Palestine, Syria and Lebanon which are discussed in more detail
below. Comparison of the Upper Pleistocene Caucasian faunal complexes
with those of Syria- Lebanon indicates that the southward migration to the
Caucasus of European forest forms (such as pine marten) followed
the Aegean route (over the Balkans and the marginal ridges of Asia
Minor) more frequently than the route over the plains of southern
Russia and Ciscaucasia.
162
168
During the Pleistocene the Caucasian Isthmus was ап ecological barrier
to the dispersion routes of the Russian Plain and southwest Asian species.
The barrier prevented the southward migration to Transcaucasia of a number
of steppe mammals of the Russian Plain, such as Siberian polecat and
suslik. The barrier also blocked the migration of animals of the upland
steppes of southwest Asia (southern-type gerbils, mole vole and other
species) to Ciscaucasia.
The Recent fauna does not reflect a series of xerothermic phases in the
Pleistocene.
At present one can speak of only one phase of development of the steppes
in the Middle Pleistocene, the postglacial dry climatic phase.
The warm (interglacial?) phases during the Pleistocene did not bring a
return of the subtropical fauna or flora from south and southwest Asia and Africa,
even to Transcaucasia. The fauna of the foothills and low plateaus remained
xerophilous and relatively thermophilous in character. Some species of the
Asia Minor and Iranian uplands moved north by way of Transcaucasia, and
animals of the Turan deserts migrated south and west through Ciscaucasia
mostly through the Manych area, but there were no new immigrations of
southern species into Transcaucasia.
The patterns and stages in the development of the Pleistocene Caucasian
fauna are better understood when the fauna is compared with the Pleistocene
faunas of the adjacent areas: the Russian Plain, the Crimea and southwest
Asia. Comparative studies can be done on the Middle and Upper Pleistocene
faunas; the data for the Lower Pleistocene and for the Pliocene- Pleistocene
transitional period are insufficient.
The total number of known species of mammals in the Middle and Upper
Pleistocene of the Russian Plain is 74. (The number is based on the
identifications of Gromov (1948), Pidoplichko (1954) and our unpublished
data for the Volga and Don areas). The faunas of the Russian Plain and of
the Caucasus have 36 species in common, which amounts to nearly 50%.
The quantitative faunal resemblance might be even higher if the alluvium
in the river valleys of the Ciscaucasian plains were to be successfully
searched for fossils.
The previously noted absence of boreal species (arctic fox, blue hare,
lemming, reindeer and musk-ox) and the presence of mountain species
(goat, chamois, sheep and a number of southwest Asian upland rodents) in
the Caucasian fauna of the Pleistocene are the main features that distinguish
it from the fauna of the Russian Plain of the same period.
Of the 58 species of Pleistocene mammals known in the Crimea (Birulya,
1930a, b; Vinogradov, 1937b; Gromova, 1935а; Gromova and Gromov,
1937; Gromov, 1948), 35 species (approximately 50%) also occur in the
Caucasus. Among the species in common are the following representatives
of plain and steppe assemblages: long-eared hedgehog, wolf, fox, corsac
fox, cave hyena, badger, Panthera leo, great jerboa, small five-toed
jerboa and mammoth.
The main difference between the faunas of the two areas is the absence
in the Caucasus of steppe species (small pika, large-toothed suslik,
Eversmann's hamster and yellow steppe lemming) and of northern species
(arctic fox, blue hare and reindeer) which are highly characteristic of the
Upper Paleolithic of the Crimea. The pronounced resemblance in other respects
of the large-mammal faunas of the Crimea and the Caucasus may indicate
169 that the Pliocene-Pleistocene faunas in both regions evolved from a common
163
stock, and even that the regions were connected in the Lower Pleistocene
by a landmass in the area presently covered by the Sea of Azov. It was the
fauna of the southern part of the Russian Plain which primarily contributed
to the faunas of the Crimea and the Caucasus.
Any discussion of the faunal relationships in the Pleistocene between
the Caucasus and the areas to the south and southwest must be based on
a sound paleontological record of the southern Black Sea coast and inner
Anatolia. However, there are no data on the Pleistocene faunas of those
regions, although the activity of Acheulean man has been traced in the
interior of the country by Pittard (1929), Pfannenstiel (1941) and Sauter
(1948).
According to Sauter, the interior of Asia Minor (Lake Tuz-Golii and
other regions) was inhabited by prehistoric man from the early stages
(Chellean-Clactonian-Acheulean) to the Mesolithic. This indication of
natural conditions favoring the maintenance of human life means that large
mammals were abundant in the Quaternary in regions bordering the Caucasus
which are today deserts and salinas.
Furon (1955) has noted that inner Turkey was less of a desert in the
Pleistocene than it is in the Recent. Lakes Tuz-Golii and Budur-Golu were
100 м deeper than their present depth. Of the Pleistocene cold phases, оп1у
the Wurm phase is recognizable.
The Pleistocene fauna of the southern parts of southwest Asia and
northeastern Africa is known mostly from the studies of Blanckenhorn
(1901, 1910, 1921-1922), Vaufrey (1931), Bate (1937), and Picard (1937).
Of the 69 species reliably identified by these authors, 22 (approximately
31%) also occur in the Caucasus. If species which are in doubt (Panthera
leo, Bos and woolly rhinoceros) are included, the resemblance is even
greater. As Picard correctly noted, there is no evidence that elk and
reindeer lived in Syria or Palestine in the Pleistocene. Nor did they live
in Egypt, contrary to Berg's uncritical quotation (1947, p.82) of
Blanckenhorn's statement on the occurrence of these northern forms. In
the study of the development of the Caucasian fauna, it is important to take
into account the occurrences of pine marten, European wildcat, Asia
Minor hamster, tiger polecat, roe deer and other forms in the
Pleistocene of Palestine. The late, postglacial stage of infiltration of
southern Asian species into the Caucasus is represented by occurrences
of jackal, striped hyena, jungle cat, kulan and goitered gazelle in the Upper
Pleistocene. The core of the Pleistocene fauna of Syria-Palestine are the
southwest Asian and north African (Mediterranean) forms, such
as civetcat, lion, true ass, warthog, hippopotamus and Sinai ibex. The
distribution ranges of these species in the Pleistocene probably did not
include areas near the Caucasus. The confirmed absence of mammoth and
the presence of the Acheulean Elephas trogontherii are important
indicators in tracing the extinction of some ''northern'' species and the
southern range extension of others (via the Aegean landmass and the
Caucasian Isthmus) in the Pleistocene.
The paleontological material of Palestine has been studied in more detail
with reference to evolution of climates and landscapes than has the material
of the Caucasus. Bate (1937) attempted to represent the climatic changes
graphically. Taking the bone-count as an index, he used several gazelle
170species as representative of а xeromorphic landscape, and the fallow deer
164
ЕРОСН$ Percentage and total numbers of bone fragments CLIMATE
AND AND
100 3 80 7 60 50 40 30 20 10 O
STAGES 10 20 30 40 50 60 70 60 90 10) Г“
Neolithic 2) Уегу агу
Mesolithic 2066 and warm
Advance of
warmth and
aridity
Slight
humidification
Aurignacian
Increasing
aridity
Warm
Humid with
Dama sharp changes
mesopotamica in the fauna
Dry and
pao Rena
warm
Mousterian D Recurrence of
dry conditions
Warm and
humid, with
occasional
aridity
a
Acheulean
Mixed fauna
(hippopota-
muses and
rhinoceroses)
Е Humid,
Acheulean tropical
FIGURE 77, Variation in number of bones of Gazella and Dama during the Anthropogene
based on food remains in caves of Mount Carmel, Palestine (excavations of Garrod and Bate,
1937)
(Dama mesopotamica) as representative of broadleaf forests and of
amore mesophytic landscape. Gazelle and fallow deer were the main food of the
ancient Carmelites inhabiting the Wadi el Mugharah and other caves.
Bate's study showedthat Dama bones predominate over gazelle bones
mainly in the Lower Acheulean, in the Upper Mousterian and in the Upper
Aurignacian. According to Bate, hippopotamus and rhinoceros lived in
Palestine at the time of the Lower Acheulean when the climate was dry
and warm, as it was inthe Middle Mousterian. The Upper Mousterian brought
a humid, pluvial climate, with an abrupt change in fauna, and the Upper
Aurignacian marked the last weak advance of humid conditions (Figure 77).
The pluvial stages in Palestine probably corresponded to mountain
glaciations of the Lesser and Greater Caucasus. Picard (1937) concluded
165
that statements of Blanckenhorn's (1910, 1921) and others that the
Pleistocene climate of Palestine was similar to the present climate of
171 Germany are erroneous. The paleontological data indicate that the climate
of Palestine was of the Mediterranean type, fairly stable and dry, from the
time of the Pliocene. :
Cooler and pluvial periods, according to Picard, are represented by
gravels possibly corresponding to the major glaciations in the north. The
main pluvial stage in Palestine correlates with the maximum, Riss,
glaciation of Europe. In addition, two poorly developed pluvials occurred
between the Acheulean and the Mesolithic. As a whole, the landscape of
Palestine was dry desert with islands of broadleaf forests in the more
humid valleys, that is, a landscape similar to the Recent.
Three localities with Pleistocene mammals are known from the region
south and southwest of the Caucasus (northwestern Iran). The mammals
occur in diluvial sediments of marly hills near the town of Maragheh and
in the Bisotun and Tamtama caves. A. Gebel's collection from the
MarAagheh vicinity (deposited in ZIN; see Brandt, 1870) contains fragmentary
bones showing the Pleistocene type of preservation: Canis lupus,
Crocuta spelaea, Rhinoceros ci: чево паз Equus
ecabatius-HrecirhremionvSst, Boersp., "OVS ch" ammon.
Paleolithic beds of the Tamtama cave, west of Lake Urmia near Rezaiyeh,
which was excavated by Coon (1951), yielded fifteen identified mammal
species, e.g., gerbil (Meriones), porcupine (Hystrix), deer (Cervus
elaphus), goitered gazelle (Gazella subgutturosa).* Bones of deer
constitute nearly 66%, and bones of horse nearly 12% of all the collected
material.
Sixteen species of mammals are known from the Upper Paleolithic and
latest beds of the Bisotun cave, near Behistun (a rock landmark) on the
Baghdad-Hamadan road. The finds of jackal (Canis aureus), panther
(Panthera pardus), pika (Ochotona sp.), goitered gazelle and other
forms are of considerable interest. The most abundant are deer (nearly
50% of the collected material); these are followed by horse (23%) and
goitered gazelle (12%). A seal tooth reported from this site was probably
introduced from the Caspian coast; beaver remains were erroneously
recorded.
This extremely interesting material indicates the northern range limits
of some southern species not found at the Binagady locality. The fauna
indicates a late Pleistocene climate and landscape in the northern part of
the Iranian plateau quite similar to the Recent. However, the abundance
of deer indicates well-developed tugai forests, which were later destroyed
by man.
The stratigraphic and geographic distribution of Pleistocene species
in the Caucasus and adjoining areas is given in Table 62.
The figures in Table 28 reflect the extent and detail of present knowledge
of the Pleistocene ''faunas'' of the Caucasus and adjacent regions. From
them it is possible to estimate the degree of faunal resemblance and the
age of faunal connections between regions. They also confirm what has
already been said of the predominantly northern and northwestern
(southwestern [sic]) influence onthe fauna of the plains and foothills of the
172 Isthmus in the Middie and Upper Pleistocene. At that time the areas of
“ Following Fraser's identification, Coon mentions the occurrence of a European beaver (Castor fiber),
which is evidently a misidentification of porcupine bones,
166
distribution of the southern (southwest Asian) and local (Caucasian) Species
either had already moved to the south, or had been reduced to isolated
relicts.
A survey of the stages and paths of development of Caucasian fauna by
zoogeographic subdivisions can be made at this point only for the Upper
Pleistocene, for which the following categories are recognizable:
1. Ciscaucasian steppes and forest-steppes, including the Ciscaucasian
plains, the Stavropol Plateau and the Taman Peninsula, from the Manych
in the north to the foot of the Bolshoi Range in the south, and to the
Apsheron Peninsula in the southeast. The fauna is characterized by
mammoth, bison, horse, cave bear, giant deer and за1са.
TABLE 28. Degree of similarity between Middle and Upper Pleistocene mammalian "faunas" of
the Caucasus and adjacent territories”
Caucasian
Isthmus
Russian Plain Crimea Southwest Asia
Number of species known from
Middle and Upper Pleistocene...
Percentage of species in common
between Middle and Upper
ЩО 55 pon od Oe oO бовос
69
* These estimates rapidly become obsolete as new localities are discovered and new collections made
available,
2. Caucasian mountain region, including the mountain system of the
Greater Caucasus. In this region mesophilous mountain-forest types of
mammals predominate: European brown and cave bear, mole vole, goat,
chamois. Also present in the eastern part of the region are a few xerophilous
species: Radde's hamster, wild goat and little suslik. These species
originated in the Lower Pleistocene and have survived to the present in
Dagestan and Kabarda. :
3. Transcaucasian foothills region, which includes the foothills and
the plains of western and eastern Transcaucasia. The eastern section of
the region is characterized by southwest Asian rodent, southern-type
rhinoceros, and horse and tur; the western is characterized by bison.
4. The region of the Lesser Caucasian plateau, characterized by an
abundance of horse, mouflon and Bos primigenius. The hot valleys
of the southern part of the region were probably inhabited by the camel and
the northwestern slopes by mountain-forest species: mole vole, goat,
chamois.
167
173 HOLOCENE LOCALITIES
174
Conditions under which animals died and formation
of deposits in the Holocene
The conditions under which animals died and their remains accumulated
were generally constant from the Pleistocene into the Holocene but for some
important changes in the later period. As the dry warm period advanced,
there was a reduction of slope erosion by running water and consequently
the deposition of diluvium and alluvium decreased, and fewer bones and
animal remains were transported to river deltas, lakes and marine bays.
Man began to affect the landscape markedly by cutting and burning trees
and changing the flow pattern of streams and, as the result of his direct
and indirect influence on the landscape, the large mammals of the plains
began to decrease rapidly in the Holocene. As another consequence, natural
deposits of animal remains became very rare.
Some cases of mass death of Recent animals from natural causes and
the modes of their deposition are described for the plains of the Caucasian
Isthmus by Kolesnikov (1950) and Vereshchagin (19514).
Mesolithic, Neolithic, Copper-, Bronze- and Iron-Age sites with bone-
bearing beds have been discovered (Piotrovskii, 1949) and these remains
of human cultures of the postglacial epoch in the Caucasus have been
thoroughly studied. The map of Paleolithic localities, shown in Figure 78,
is after Krupnov.
The scale and the location of human settlements changed radically in
the Neolithic. Settlements were located on the shores of lakes and on river-
banks, andcave dwellings remained only as relicts of the Paleolithic until
they reached a new stage of development during the feudal wars of the Middle
Ages.
Because there was little tectonic activity in the Holocene, marine,
deltaic and lacustrine deposits remained undisturbed, as they had during the
Tertiary and the Pleistocene, and consequently they lie unexposed today.
The possibilities of finding bones in relatively young deposits in diluvium,
small niches and caves are considerably greater. Such accumulations of
bones resulted from temporary settlements and predator activities,
particularly those of owls and eagle owls. ©
Bone fragments similar to the Paleolithic constitute the food remains
in the deposits of Holocene settlements. However, many more bone and
horn implements were found from the Holocene than fromthe Paleolithic.
Occurrences of bones gnawed by domestic dogs are highly characteristic
of the Holocene, but phalanges and metapodials from which bone marrow
was extracted were no longer as thoroughly broken as in the Paleolithic.
Beginning with the Eneolithic, certain bone accumulations inthe Caucasus
which contain complete skulls and skeletons of both wild and domestic
mammals give evidence of being ritual burials. The religious-cultural
customs of several Caucasian mountain tribes account for peculiar
ceremonial accumulations of domestic and wild animal skv’.s in the last .
few centuries of our era.
The discussion which follows, organized into geomorphological regions,
treats all these types of deposits, some of which have been studied to a
greater and some to a lesser extent.
168
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169
175
Western Ciscaucasia'iand the south
Rue Sane Вата
The establishment in the Holocene of firm land connections between the
Ciscaucagian and south Russian plains enabled mammals to migrate freely
from one region to another.
Investigators studying Pleistocene fauna in the lower Don region
uncovered abundant mammal remains from the Holocene. These bones
were washed out of floodplain sediments and redeposited on sandy-gravelly
spits. Archaeological excavations of settlements and mounds in this same
area which were carried out by the Institute of Material Culture, AN
S.S.S.R., also yielded large collections of bones.
Particularly valuable bone material was collected by Gol'msten during
the excavation of mud-hut settlements on the Tsaritsa River dating from
the fifteenth to tenth centuries B.C., and during the excavations of the
Khazar fortress of Belaya Vezha (Sarkel) near the village of Tsimlyanskaya
(Artamonov, 1952).
Eleven species of domestic and 23 svecies of wild animals have been
identified from the more than 30,000 bones collected in these excavations
on the lower Don, mainly at Belaya Vezha. *
The faunal complex of the steppes and plain forests is still pronounced
in the wild mammal fauna of this period. The assemblage of game animals
includes 16 species, ranging from corsac fox, steppe polecat and beaver
to bear, elk and bison. The materials indicate that even in the Middle Ages
(700 to 1300, A.D.) the ranges of forest animals (taiga and forest-steppe
zones) extended along the river valleys to the Ciscaucasian plains, where
in the thickets of the floodplains of the Yeya and Kuban rivers they joined the
Caucasian ranges of the same animals.
TABLE 29, Mammalian species and number of bones from Early Bronze Age mud huts near Gelendzhik**
Number ЗА
of bones Saas
2
55
Number
one of bones
Domestic
HIOISE РО СО ооо ОНО ЗС РЕ ПУХ а сома
Ради © тала оды. Depirs den ropiaS Ws Fes + eters 6 oe
рат ется ну ль, се 396 Died pihwmirs? Чери. лье fe
Small cattle, primarily sheep .,... 106 Sills sSi@niayfval «he. о бов. abe
а ЕЕ 559 ое: ВАН, "АН
Аа Ми Г.
6 pe езде" 0) oN) Фе
The main archaeological monuments within Ciscaucasia proper are
Neolithic settlements, Scythian mounds and cultural layers many meters
thick of ancient Greek colonies and of the so-called Bosphorus kingdom
(Rostovtsev, 1918; Kallistov, 1949) on the Taman Peninsula and on the
* This material is being published separately.
** This and other similar tables were compiled from the author's unpublished data.
170
Kuban and Trans-Kuban plains. The Gelendzhik settlement is among the
oldest settlements of the region.
In 1952 Akhanov collected many bones from pits in the Upper Neolithic-
Lower Bronze Age mud-hut settlement on the shore of the Gelendzhik bay.
The material consists of mottled and yellowed diaphysal and vertebral
fragments which, whentapped, emit alight, ringing sound like calcined bones.
Most of the bones are those of large cattle and dolphins; those left by man
were later gnawed by some small predators.
The wild animals are represented by deer, bear, lynx and hare (Table 29).
Individual bones of fish, shells of marine mollusks and edible snails were
also collected at the site.
The wild species at the Gelendzhik settlement indicate that sea game
was important to the ancient Tamanians, or at least that they consumed
the bodies of dolphins washed ashore by the sea. Forest game species
(lynx, boar, deer) were also abundant in the vicinity.
Thick bone-bearing beds, usually no more than 2-3 m and occasionally
as much as 10m thick, occur on the shores of the Taman Bay, at the sites
of the ancient towns ofCepi, Phanagoria and Taman. The beds are being
176 eroded by the sea because of the subsidence of the coast in historical times.
Bones of domestic animals, sturgeon-like fishes and characteristic layers
of purple Mytilus shells comprise the food remains which are plentiful
at these sites. Bones of wild animals rarely occur in these beds, which
confirms the accepted concept of a high degree of cultural development in
this region at that time (Figure 79).
FIGURE 79, Exposures of cultural layers of Phanagoria on south shore of Taman Bay
Photograph by author, 1952
171
Bie
178
TABLE 30, Species and number of bones of domestic mammals from Taman towns
10
Species
Зо 06 бое зе дея del Малое леща ide heen Ва (6 6) un eae
а eon fore Toke be We Rel ее ее ime cist ее
ELOISE? С ts Me ants НЫ к tiene. hn aye, es 11 18 13
Е, Ель РСЯ He 6 fey?
TAREE Callen, whee, TTT iy а нь вы ete ae о 68 27
SHEGP BALLER yaaa Paes PLS eR Ie 15 34 32
а aye: © 9 einoniewel je) р ei weve, Юве ии о shes Зы Эра
о e te! tee С, ОВ
Table 30 lists bone fragments collected from several hundred meters
of cultural layers from the three towns.
From the incidence of identified domestic species it appears that the
landscape of the Taman Peninsula in the second half of the first millennium
B.C., on the whole, was similar to the Recent. The nomadic herds of the
Scythians and the local animal husbandry of the Greeks must have strongly
depleted the vegetation cover of the peninsulaatthat time, Large species
decreased and the remainder of the animal assemblage probably consisted,
as it does in the present, of European hedgehog, fox, hare, field mouse,
house mouse, common vole and mole rat.
More complete and interesting data are available for the ancient site of
Semibratnoe near the Cossack village of Varenikovskaya, east of the Kuban
River mouth. Anfimov's excavations in 1938-1939 and in 1951-1952 showed
that a small settlement occupied the site as early as the fifth century B.C.
A town of considerable size developed in the third to second centuries
B.C., whichwas later destroyed and abandoned. A small settlement of
hunter -fishermen occupied the site in the first century A.D. The bone
material, collected in stratigraphic sequence, completely confirms the
conclusions drawn from study of the material culture. Particularly
Significant are the large numbers of bones of wild species occurring in the
fifth century B.C. and again in the first century A.D. (Table 31).
Some bird bones were also collected in the beds of this town: domestic
chicken, wild duck and stork. The collections also include numerous bones
of sturgeon, stellate sturgeon and cyprinid (mostly wild carp).
Ancient animal husbandry was evidently not highly developed in the
Bosphorus. Dogs were uniformly small, about the size of a spitz, and were
probably used only as watchdogs. The bones of goats, sheep and swine
are light, which indicates poor feeding. Cows were of the European short-
hornedtype, somewhat larger than northern ''peat'' cattle. The occurrence
of bones of otter, boar and deer may indicate that reed-grown swamps
covered the area, in which there were probably no large forests.
Bones from excavated Scythian mounds in the Maikop area, dating from
the middle of the first millennium B.C., are known only from Vselovskii's
drawings (1901). He depicted many tens and hundreds of skeletons of
horses which had been killed during funeral ceremonies for tribal chieftains.
172
TABLE 31. Mammalian species and number of bones from the ancient town of Semibratnoe
ln) present Before Present Era a
р Ега 3
Species ae)
Gs}
I I II Ill с Е
Domestic
DO Gel ен ae ay ecg 46/32 25/16 48/31 99
FIGTS CAMERA он Зо НЙ 7/5 10/6 102/52 33/18 74/32 | 142/75 188
оао, ai aia Aare i pa Ha ek 15/8 5/4 64/44 36/16 | 101/42 | 103/70 184
LATS CBWE 5 5 4 ove.6 prove oo 6 on 39/14 13/5 153/67 87/37 215/53 | 303/183 309
SITES Pee ets cee heme. etc Pn ee ee sae, 9/6 13/12 72/43 27/11 98/37 | 152/72 181
Goan oe He Re aan ae NUL es 17/8 32/17 63/28 56
Small cattle (no closer
ПОМИ ЕМЕ о )) 5560000007 во
ДРОВА so Aen a boo old
Wild
(GAIUS ell U Sons омвибанно овен ©
MENGS. THENCE 5 S% 516 ое
аа ета» 9. Зоо, 5
РЗ CULOPEBCWS 4555050006
SMS БОНО ees eee 25/7 2/1 4/2 3/3 2/2
Cenyms ClapmMWS 545 66s bo 24/7 12/8 6/6 4/3 6/6
Capreolus capreolus..... = = 1 =
il ae
SUDTOT pos0008 Fae aca ican 80
СЕ ipreneiak Silber allies 1,101
Note, Slash separates number of bones from number of individuals.
TABLE 32. Mammalian species and number of bones from mounds near Ust-Labinskaya
Present
Before Present Fra
Species
individuals
Dog Duc onokor.a 020.0 oLoke 3
HOMIES 5 о bb 900000004 we
О о а 6 13
БИТЬ CANIS оо ороовос 18
SINE CDs: оке НО 5
COL ево orotote ss о 3
Small cattle (no
closer identification), .
ооо
Note, Slash separates number of bones from number of individuals,
3:
eg
Excavations on a limited scale carried out by Petrovskii and Anfimov
(1937) in the towns and burial grounds of the Kuban area east of Krasnodar
uncovered bones of horses and large and small cattle from layers of the
fourth century В.С. to the third century A.D.
Pieces and amulets carved from large deer antlers and bear tusks which
were collected from these burial grounds are housed in the Krasnodar
Museum.
We identified only eight species of domestic animals among the 337 bones
from the burial grounds excavated by Anfimov near the Cossack village of
Ust-Labinskaya. The animals were sacrificed during funerals and buried
with the dead as food supply (Table 32).
The dogs in this region were about the size of a small husky. Swine,
cows, sheep and goats were also quite small.
The ancient site of Elizavetovskoe and mounds on the left bank of the Kuban,
13 km west of Krasnodar, were excavated by Gorodtsov (1935, \1936) in
1934-1935. From layers of the early centuries of the present era,
Gorodtsov recorded ''many bones of cows of two varieties, small pig, sheep,
goat (?) and three varieties of dog. Bones of wild animals are very rare;
only antlers of red deer, placed in the grave with the dead, were found."'
Artamonov (1937) collected remains of the following animals from Late
Bronze Age mounds on the steppes of the northern bank of the Manych
(near the Manych canal and the village of Spornyi):
Domestic Wild
Horse Lepus europaeus
Pig Marmota bobac
Large cattle
Sheep
TABLE 33, Mammalian species and number of bones from hills on the south bank of the Manych
Species Number of bones Number of individuals
Domestic
DGG TS ius оао: - ayy os eee omens leuate ite 16 1
PLOIS GP Оки Нее rae cco НИИ не ББ 12 2
Targe cattle. Sats cutie aie te eee te ole wie, ла 4 2
Sheep) i A Re les J A eed 288 Pe Ta 5
И о FY ek: в Дрю 103 10
Wild
Vulpes ‘wnlpes red... ob а. 21 38 3
сама, Пири ste cme eval eee ees sie eee se 1 i
Mie les mie'les леса ее 1 1
Tepisretrope Gus ease te 7 pean nme aie n er 2 1
SUD BOG ИИ ПРЕ у оон aon tae 42 6
RAR cg tee щи целы vitae 145 16
174
180
Other species are known from the southern bank of the Manych, near
the village of Veselyi; they occur in mounds of the middle of the first
millennium B.C. (Table 33). Had marmot bones been found at this site,
it would be of great paleogeographic interest, since marmots have not been
documented on the Ciscaucasian plains by twentieth-century zoologists.
Domestic animals were probably sacrificed in rituals. The bones of fox,
wolf, badger and even hare, however, which were collected from the talus
of the mound, are the remains of animals which died in their holes.
The foothill regions and river canyons of northwestern Ciscaucasia are
paleontologically unkncwn. Interesting data can probably be obtained from a
study of the bones in the ancient Alani settlements, which are preserved
in ruins in the Laba, Belaya and Zelenchuk river gorges and in the old pits
in asphalt crusts near Khadyzhensk (Vinda, 1910).
The only bones known from this region are the skeleton of Microtus
(Chionomys) roberti —from a 14th-century burial chamber in the
gorge of the Malaya Laba River, near the village of Andryukovskaya — and
a horn of a Caucasianelk (Alces alces caucasicus), collected in
1949 in the gravels of the Urup River near the village of Otradnaya.
@Genitwal Cis cawicasia
At present the Stavropol Plateau and Pyatigor'e are peculiar meadow-
forest ''mesophytic islands'' in the steppe plains. As such, they have
retained to the present a number of mesophilous species of mammals which
migrated to this region during the cold phases of the Pleistocene. The
migrations came from the north, from the south and from the Greater
Caucasus.
Deer and lynx which have completely disappeared from the relict forests
of the Stavropol area are known from published records (Dinnik, 1914а)
and from material in the Stavropol Museum to have inhabited the region
until recently.
Small mammals are among the species which have disappeared, or are
about to. The length of time that the little suslik was isolated in the inner
valleys on the northern slopes of the Greater Caucasus is a particularly
interesting paleontological problem.
Toward its solution, bone material from the layers of the ancient towns
has been studied and special reconnaissance excavations have been carried
out in caves and under overhangs on the margins of the Stavropol Plateau,
in laccoliths of Pyatigor'e and in cliffs of the river canyons cutting the
Kabarda Plain.
The region of the Kabarda Plain was well populated by man in the Lower
Holocene, as shown by occurrences of Neolithic tools, numerous mounds
and remains of ancient settlements. The Eneolithic burials at Nal'chik,
excavated by Miller in 1929, contained ornamentations made of teeth of
domestic and wild animals: cat, fox, boar, red deer, goat, sheep and a
very large Bos, either domestic or wild.
Similar ornamentations from excavations by Kruglov, Piotrovskii and
Podgaetskii (1941) were found in the Neolithic burials of the Nal'chik mound,
175
near the ancient site of Agubekovskoe. The ornamentations consisted of
teeth of boar, deer, fox, bear, and Bos.
The Degen-Kovalevskii (1935) excavations of the 6th—8th-century
settlement near the village of Zeyukovo (53 km from Nal'chik) found the
Alani food remains to consist mostly of bones of domestic animals (Table 34).
The bull from this locality was small, as shown by its bones, and
undoubtedly belonged to a domesticated variety. Characteristic of the
locality is the first occurrence of domestic pig and of a very primitive
variety of goat, with horns curved similarly to the horns of Capra
prisca.
Kistyakovskii (1935) recorded remains of 15 species of extant rodents
in the pellets of predatory birds of the steppes to the north and east
of the Stavropol Plateau, and reported their occurrence in the same area
of distribution as at present.
Some bones of rodents and insectivores were collected by Lyutyi (1940)
181 from eagle owl pellets on the western slopes of the plateau, near
Stavropol. His data did not contain any new zoogeographical information.
TABLE 34, Species and number of bones of mammals from Alani settlement near Zeyukovo
Species Number of bones Number of individuals
Domestic
О а беби Geeks оао 2 1
И Ор А 8 2
о ре Е 15 3
ато ВС а ее ое о о ses 29 3
Sriall саме"? Jt: FES APA Ve eS = 10 2
а ое ро а о aun enol ay ieee 64 1
Wild
1
Rordionaa(aush-oved- sabi 12
In the fall of 1950 we made a paleontological reconnaissance survey of
the western part of the limestone cliff of Mount Strizhament on the
southern margin of the Stavropol Plateau. The presence of forest, springs
and "а sea of rocks'' indicated a likelihood of finding bones of forest and
steppe animals, left by ancient hunters and cave predators, in this area.
In four small caves only individual bones of sheep and cows, brought
by shepherds and wolves, were collected. The unstable ceilings and big
blocks of limestone on the floors prevented a more detailed survey.
A considerable number of small-mammal bones brought by eagle owls
during the last century were collected in one of the caves on the cliff.
The results of the identification of the bones are given in Table 35.
The most interesting find was the remains of a pine vole, which is not
recorded in the literature as occurring in this region, although the species
176
commonly occurs nowadays оп the meadows of the plateau. This
occurrence of the species, Separated from its main area of distribution in
the Caucasus, is biogeographical evidence that the climate in the past was
more humid than in the present.
Pyatigor'e. Studies were done on isolated elevations in the steppes
of Pyatigor'e and compared with the Stavropol Plateau. Particular attention
was devoted to the permafrost of Mount Razvalka, 5 km north of
Zheleznovodsk, and to the gorge of the Berezovka rivulet south of
Kislovodsk.
Reconnaissance excavations were carried out on the eastern slope of
Mount Razvalka near the Selitryanaya cave. At the site, dense broadleaf
forest surrounds two picturesque trachytic rocks, weathered by wind
erosion. A small dry cave, formed at the junction of three vertical joints,
182 is located under the southern rock. Over the last millennia, deposits
accumulated to a depth of 8-9 m under the entrance to the cave through
erosion of the upper cliff platforms and the rock face, and as the ancient
dwellers cleared the cave of rock fragments. The site of the debris was
probably used as a resting place for cattle, since the bones of small and
large mammals and fragments of pottery occur in layers at the foot of the
cliff and in the cone of the cave debris.
TABLE 35, Mammalian species and number of bones from pellets of eagle owls on Mount Strizhament
Number of bones
Species Number of individuals
ООО МИХ oo боозоровоновов
ПОЛА ССИ CUPOMACWS сооборовосовоооо
WANDA CAWEBSICA 5 оорообововоовьоеьв
Mustela nivalis caucasica
IMIS VEL INS AGM BG oe oo e ood ono OOd
Шри CUROPABSCWS соборов осбовосроос
СТАС ФТС ШиИВ - оо одосвоочооовоес
IMCSOCRICCTUS AMRATHS оообовоовососое 34
CHiCSHOINS мата ПО 55 500000000c0% 29
MR ОТО ЯЗ 25 o50b0a00000000005
МИ, MIAVORL Soond0000000000000000D 0008
рада впору ИЕ ИИ ooo 5r0000c0n 06
ADOGESWAOS БУТИ вБообочбоовоосос
MICROWAVE ОМИ сроровобовыыоовосов
MDS ИОВ ОВ © оо о обо oo ob во о ano eo
CN = не ю
нон 6 - ©
oe ee ee ооо
oo ee ee ee ee ew ee ee
Our trial trench, 1 m wide and 3 m long, was dug 1.5-1.9 m deep
into the talus in front of the cave entrance. The collections
consisted mostly of simple black pottery, made with broken shells, of
ancient Alanitype. The pottery fromthe deeper layers is ornamented with
pitted and striated markings, characteristic of the Bronze Age. An obsidian
fragment, a fragment of a stone hammer and a small whetstone made of
black slate were found in the upper 60-cm-thick layer.
177
TABLE 36. Mammalian species and number of bones at the Selitryanaya cave in Pyatigor'e
3)
Dept п
bones
рЕНЕтх
Domestic
И ее ао Oe qari ole 2
ор cloned Maumee Conant iene 19
ие 58
Татр ts Geer tate naa ase 216
Sheep’. SG DTS 175
я ВЕ С PAE) Se 34
Small cattle (no closer identification) 176
Step portale YS ae oe eee 680
Wild
Нерест ‘атс. 2
Erinaceus europaeus...... 2
Ура cia Nicasitcal к eae ats 3
Vulpes val pes ncaa 1
МО Зе COSa о 1
вета ма ео 3
Medes-mielesse sem arent 4
Putorius eversmanni...... 2
Wowie lia: Sainimatica buen, i
ме ме со ri i
Lepus europaeus... fas ss ‘ 19
о te ее 15
оО обо bre 197
Mesocricetus auratus ..... 183
Cricetulus migratorius.... 17
Arvicola terrestris д... 41
Microtius amvalis: smb. oo 18
ME Artal орех sa 2
Ratti: помех CUS denelsenertemsns 2
Apodemus sylvaticus ..... 4
Мс аи се oh nents о ОИС 1
Spalax microphthalmus, aL
ОЗЕРА ое ово ete eis 12
Gervils) ela mins и Sisal 5
Gapreolus wapreolus sn. 6
Зидан о ТЕ 543
yO Ball серое ин 1,223
Fragments of large-mammal bones are typically found as food remains.
The burial also contains rare fragmentary implements made of deer antlers.
Remains of small animals are represented by bones from the pellets
of birds of prey, mostly eagle owls which habitually rested on the rocks
above the cave. The locality is a rare and valuable example of a deposit
formed by the three processes of bone accumulation: remains of food
178
transported by birds, remains of game transported by hunters who used
the cave for shelter, and remains of food left from ritual tribal feasts or
transported by ancient shepherds.
Nearly 1,500 fragments of domestic and game animal bones were
collected in stratigraphic sequence, all showing an approximately uniform
mode of preservation. In small fracture, they are yellow or white in color,
and very few show any signs of fossilization.
184 It can be estimated that the layer accumulated over the last 2,000-
2,500 years.
FIGURE 80, Cave inhabited by eagle owl on Mount Lysaya.
On top are bodies of seven hedgehogs and one corncrake
Photograph by author, 1952
Six species of domestic and 26 species of wild animals have been ,
identified in the collection of bone material (Table 36).
Remains of birds, toads, frogs andshells of Clausilia and Helix
were collected at the same locality.
179
185
At the time of the ancient Alani settlement most of the Razvalka slope
was probably deforested and man's occupation of the site probably drove
away the eagle owls and other predators. As a consequence of their
disappearance from the area at that time, the deeper layers of the deposit
show a Sharp decrease in the number of rodent bones. In the main, the
deeper layers reflect man's activities, and it is only in the last quarter of
our era, when Pyatigor'e and Razvalka again became covered with forests,
that most of the wild-animal bones, particularly those transported by eagle
owls, were accumulated. Large forest game inhabited the area at this later
period: bear, boar, deer and roe deer. Old records show that deer
disappeared from the region at the beginning of this century. Bears, roe
deer and boars occasionally enter the forests of Pyatigor'e in our times,
although they do not live there permanently. During the same late period,
the last four to five centuries, the food of the eagle owl consisted of small
species characteristic of the laccolith region in the present: three hamster
species and water, pine and common voles. Bones of snow voles
(Chionomys gud), hamsters and water rats were collected in rock
crevices over the stone-covered permafrost area.
A few specimens of snow vole were collected in the rocks of the ''century-
old freezer'' of Mount Razvalka, at altitudes of 400-500 m above sea level.
The nearest that the species has been found elsewhere is a site 3 km south
of Kislovodsk in the gorge of the Berezovka rivulet, which is 50-55 km away
from Mount Razvalka. Thus the occurrence of the snow vole at Mount
Razvalka indicates that during a cold period in the past, probably at the time
of maximum glaciation, the mountain animals migrated to these minor
elevations in Pyatigor'e.
FIGURE 81, Crevice in limestones on western wall of Berezovka rivulet gorge near Kislovodsk — day resting
site of eagle owl
Photograph by author, 1954
180
TABLE 37, Mammalian species and number of bones from pellets of eagle owls near Kislovodsk
Number of Number of
Specie Species
А individuals* р individuals*
сл
Noon FP FN FP
ARWVLOCOGOLA сот ороборовб
IMGECRO TMS ЕМУ ING бобов овею
SOrSk AVAMEWS особовововоо
WAMI92 COSECH ор ъомовосовос
MUSESIA mMiV@INS орооыадовов |) — о iG! боббообо Alo вообоовов
APOCE MNS ЗВ ТОМЬ 555000
MIMS мии 5556065005000
Са МЮ MOnVYEENIEWS 5555000500
Spalax microphthalmus
LSpwUs, ИГОРНОМ сооосоообь
DywOiwa ws munee@tMla oo 555600 оо
CrueSrws CUUG SUMS odo ood v oe
* Number of individuals given is based on number of mandibles.
Plentiful remains of insectivores and small carnivores in the pellets
of eagle owls were collected in the laccolith farther northeast under the
rocky overhangs of Mount Lysaya (Figure 80) and in the gorge of the
Berezovka rivulet, south of Kislovodsk (Figure 81).
186 At the latter locality the material is older and more abundant. It was
obtained from a half-meter-thick layer of Holocene dust deposit under a
vertical joint in the Lower Cretaceous limestone cliff. This resting place
must have been abandoned by the eagle owl not less than a century ago,
since it is located only 1 km fromthe southern outskirts of Kislovodsk.
The bones accumulated during several centuries of the second half of ourera.
A total of 1,582 bones was collected. Their preservation is not uniform.
The bones from the lower layer are coffee-colored with dendrites; bones
from the upper layer are yellow.
The species composition of the collected fauna indicates that the
landscape at the time of accumulation of the bones was very similar to the
present (Table 37 and Figure 82).
Nineteen specimens of amphibians and 305 bones of birds were collected
at the same site. The assemblage of relatively xerophilous steppe species
is quite prominent, as at Mount Razvalka: hamster, polecat, mole rat.
187 This assemblage never occurs in complete composition this far into the
mountains. The moist meadows and rock-covered areas of the Berezovka
gorge account for the large numbers of water rat and the occurrence of
snow vole.
The hamsters and mole rats in this region were the precursors of the
relatively xerophilous assemblage which migrated in postglacial time from
the dry Ciscaucasian plainsto the north. This advance overlapped by tens
of kilometers the northernmost occurrences of the forest and alpine species
(pine and snow voles, forest dormouse, fat dormouse) which remained
in mesophytic sections of the laccoliths and in the river gorges after the
recession of the colder, more humid period. The uppermost limits of
distribution of this assemblage are the moist meadows of the Pastbishchnoi
ridge and the lower reaches of narrow gorges cut by rivers through the
ridge (Vereshchagin, 1953а).
181
(186)
FIGURE 82, Mammalian bones from pellets of eagle owls in the Berezovka gorge
1 —humerus of Talpa caucasica; 2—skull of Mustela nivalis; 3 — pelvis of Lepus
europaeus; 4—jaw of Rattus norvegicus; 5,6—skull of Cricetus cricetus; 7,8 —skull
of Mesocricetus auratus nigriculus; 9, 10 —skullof Arvicola terrestris
In the western part of the region, a similar picture exists, although at
a lower altitude, on the right bank of the meridional section of the Kuban
River.
The following conclusions can be drawn from the study of Holocene
animal remains in the foothills of central Ciscaucasia:
1. The intensive exploitation of the region by man from the Neolithic
to the present resulted in a diminution of the wooded area and the
disappearance of large animals (bison, deer and, later, roe deer and boar).
2. The presence of mesophilous relicts of the cold (glacial) epoch has
been established. The relicts survived in the steppe on the Pyatigor'e
laccoliths and on the Stavropol Plateau. This fauna includes the snow and
pine voles.
182
188
3. А dominant assemblage of relatively xerophilous species of rodents
(hamster, mole vole, mole rat), characteristic of open areas, existed in
the foothills of the Kislovodsk area and further south during the last
millennium, and possibly during the entire present era. During this period
the little suslik did not live in the Pyatigor'e area. The migration of this
species in the Holocene to the longitudinal valleys of the range can be
established only after its remains have been found in layers deeper than
those excavated. Such an occurrence might prove the existence ofa
postglacial climatic phase drier and warmer than the present climate.
Eastern Ciscaucasia
The steppes of the eastern Manych area are of interest to the
paleozoologist because of the occurrence of a microlithic culture, usually
identified as Late Neolithic. Tribal sites containing microlithic material
occur in the southern Ukrainian and Crimean steppes, along the northern
coast of the Caspian and in the border scarps of the Ust'-Urt, and deep
into Soviet Central Asia. The most permanent settlements of these tribes
were located in the Berovskie hills, surrounded by elms and rich in fish
and boar.
In this region, remains of material culture and fragments of animal
bones occur in wind-blown depressions, on wind-eroded mounds and in
sandy hills. Fragmentary bones of giant mole rat (Spalax giganteus),
Saiga and sheep occur at sites such as these in the steppes of the Kuma
area; the great gerbil (Rhombomys opimus) occurs in blackland sands.
Between the lower reaches ofthe Kuma andthe Volga, the archaegqlogist Sinitsyn
discovered numerous bones of animals of many varieties around the remains
of ancient campfires. He also discovered many settlements and burial
grounds of the Bronze and Middle Ages.
Remains of a large bull with horns similar to those of Bos
primigenius from the pre-Scythian Tri Brata mounds near Stepnoi
were photographed by Sinitsyn (1948, p.151).
He dated the burials end of the second to beginning of the first millennium
B.C. The culture, i.e., Transcaucasia, is similar to that at Trialet
(Kuftin, 1941).
Similar and younger monuments of the pre-Scythian and Scythian epochs
have been traced by Krupnov (1947) in the steppes between the Terek and
the Kuma. Osteological material from this region would greatly clarify
the notion proposed by Krupnov that there were considerable changes in
the landscape during the last millennium B.C., and would probably also
improve our concepts of the history of the ranges of central Asian desert
animals.
Farther south, on the northeastern slopes of the Dagestan foothills, bone
material is known from settlements and mounds of the Bronze Age.
Primitive archaeological excavations were carried out by Russov (1879) and
Tsilossani (1879) in the region of Derbent, Kayakent and Deshlagar. Their
excavation log indicates ''bones of rodents,''''bones of animals, '' ''bones
of large and small domestic animals,'' etc., occurring in caves,
mausoleums and burial mounds. The ZIN collections contain bone materials,
183
obtained by Kruglov (1946a) in excavations in 1941 from layers of a
settlement of the second half of the second millennium B.C. near the villages
of Kayakent and Dzhemikent. The bone fragments from these layers
are light brown in color and they clink slightly. Bones of boars tend to
absorb water as do Upper Pleistocene bones. Identifications and
counts are given for these fragments in Table 38.
TABLE 38, Mammalian species and number of bones from Bronze Age settlement near Dzhemikent village
Number of bones
Species Number of individuals
Domestic
DIO en G1, о ве...
ЕЮ Бо бое auc Ovo с а diigo. 60d. 0.6 topo aatan
StS 07 We Е: сос 60
PhoG@a C:asip Tare ss <1. Be. а...
ие Боло fl, alate Rep ecko
Уцир аз уе Re iete a sence oh ed.
Бам MGM OM Sp of bs ey ees An hs ao Soe №
ИВО о О И wea awede igs Gheds
ee ee ee ны
ооо ооо
Зо се. в. © в я ©. р би са « 0 6
189 The occurrences of seal, goitered gazelle and kulan in the region are
noteworthy (Figure 83).
The reconnaissance excavations of 1954 carried out under the picturesque
cliffs of Kapchugai west of Makhachkala by the archaeologists of the Dagestan
Branch of the AN SSSR revealed layers which have been dated late
second to early first millennium B.C. The following bones have been
identified by us from these layers: horses (3), large cattle (12), sheep (14)
and red deer (Cervus elaphus) (12). These numbers indicate both the
abundance of deer and the importance of game hunting in the economy of
the ancient shepherd tribe. The cliffs of Chokrak sandstones at the
site are covered with line drawings of arimals, made only a few centuries
ago. Panther, horse, deer and saiga can be recognized in some drawings.
Remains of Iron Age animals are known from excavations of ancient
settlements in the valleys of the Sunzha River and its tributaries.
Krupnov (1948) has published the results of excavations of a settlement
of Scythian time (second half of the first millennium B.C.), near the village
of Alkhaste on the left bank of the Assa River (Table 39).
He also reports (1949) a similar ratio of domestic species
distribution for the Assa River valley settlements of Nesterovo and Lugovoe
(sixth to fifth centuries B.C.; identifications by Tsalkin). At the Lugovoe
184
locality a horn stemof a saiga and individual bones of а roe deer have been
identified among the numerous bones of domestic animals.
FIGURE 83, Mammalian bones from the Bronze Age settlement near Dzhemikent
1 — jaw of Vulpes vulpes; 2—tarsus of Ursus arctos; 3-5 —first phalanx, hoof and astragalus
of first phalanx and tibia of Equus hemionus; 6-9—horn, head of femur, first phalanx and tibia
of Gazella subgutturosa
(190)
TABLE 39. Mammalian species and number of bones from Scythian epoch settlement near Alkhaste village
Species * Number of bones Number of individuals
Domestic
Doe ce Е : Holo n Эа О: 6 3
BIGKS® соб Sy SLND EE SLD pee Heel Mone eee wel octet wets 21 6
И О ес on Mos инь: 175 15
Largereatrle tiie! 3.20 ВЫ: мВ 101 10
Small cattle (no closer identification) ......... 54 8
АИ во бою aac aa ee isha аа 357 42
Wild
Vulpes vulpes ...... ERROR TS Athens ee 1 1
GNSS ще ети сети зм О А аа, 5 1
Сетмз fe lapis) J oe hee pele. AOE 2 1
Сархео 1 CApEGOllG Bbesceaduomoesuac 6 3
i ey eas 9 oe
TOA Lee es Sas A ee eee ate. Sates 371 48
* М.А. Sugrobov's identifications, Bones of bear were probably included with domestic pig.
191
TABLE 40, Mammalian species and number of bones from settlements and burial grounds in the
Sunzha valley
Species Number of bones Number of individuals
Domestic
DOG, ве о 102 ees 6s) 6 АЕ 5 о 75 4
Gate ТЕ... „ба oy oes .- SOS 5.0 а 5 1
Ногсе’ сы...’ М О ый. «о. 55 3
Р сое. с, о. НХ « - 32 10
Large ея е .,. 5.0% ele sss oe + 6 м. > 64 7
Sheep SRS уно saers sas оО ВЕ ме 145 12
Би ше 00$ ооо Е. il 1
Cexrvisme lapiwis’ <p en. и 9,
os в « № № ota winnie ее
Gin) (a) ‘eine, ооо (hi erie) 8 ie
In the eastern part of the Sunzha valley, excavations of settlements and
burials of the second to third centuries A.D. were carried out in 1938 by
Kruglov near the villages of Isti-Su, Alkhan-Kala and Khoro-Chai.
The bones from these excavations, identified by the author (Table 40) are heavy
and stained in places with humus. There are only a few remains of wild
animals. However, the new evidence of past occurrences of steppe species
of large mammals in the foothills of the eastern Caucasus is of considerable
interest.
In 1952 we made a reconnaissance search for species of smaller
mammals in the cavities and talus of the Sarmatian limestones above
Makhachkala and in the sandstone cavities near Kapchugai. Large numbers
of bones were collected: long-eared hedgehog (He miechinus auritus),
hare (Lepus europaeus), little suslik (Citellus pygmaeus),
hamster (Mesocricetus auratus nigriculus) and steppe vole
(Microtus socialis). All the bones were collected in the pellets of eagle
owls and flying predators which settled to rest on the cliffs.
Further paleontological studies of Holocene burials, which take into
account the earlier archaeological data (Komarov, 1879; Shtein, 1879;
Iessen, 1935), will undoubtedly fill in the detail of the general picture we
have of postglacial steppe species of ungulate mammals in the region.
They will also add to an understanding of the decrease in numbers and
deterioration of composition of the fauna toward the present.
186
193
Томе ада те vist Of) timelgnion t hemn 51 оре
ое‘ Greater @Camecacwa nw andy юоев`Базесваи
The longitudinal tectonic valleys, bounded in the north by the
Pastbishchnoi and Skalistyi ridges, are well developed east of Elburz. Like
woodless inner Dagestan, the valleys attracted post-Paleolithic tribes as
inhabitants because of their natural shelters and dry, relatively warm
climate. Mesolithic sites have recently been discovered in the valleys and
in inner Dagestan. Some Bronze Age tribes left behind them remarkable
burial monuments: mounds, underground burial sites and catacombs for
mass burials. The sites of their settlements are represented by fairly
thick layers of the ''Kobanian culture’ (Uvarova, 1900, 1902).
The medieval Alanis built numerous fortresses andwartowers, fortified
caves and individual trenches in the vertical, rocky cliffs. Their
descendants, the Ossetians, built peculiar collections of hoofed-mammal
skulls in ritual places called ''dzuars. "'
Many bones of rodents from the pellets of eagle owls accumulated in
natural niches and ruins of the fortresses.
We studied all these types of deposits, withthe exception of the Kobanian
burial grounds and settlements.
Mesolithic sites in the Baksan and Avar Koisu gorges
In 1955, Zamyatnin discovered and studied diluvial sediments, many
meters thick, containing Mesolithic flint tools, on the right slope of the
Baksanravine at the head of the first gorge near the village of Byllym
(Figure 84). Excavations to a depth of 6.85 m at the Sosruko overhang
revealed a sequence of Middle Ages, Early Iron Age and five Mesolithic
layers with characteristic flint tools of the transitional epoch.
Most of the bones taken from every layer were broken into small
fragments. The loss of organic matter in the bones is in its initial stage.
Identifications of the collected bone fragments are given in Table 41 and
Figure 85.
In addition to the species listed, about ten bones of partridge, short-
eared owl, griffon, vulture and small Passeriformes were collected
from the Mesolithic layers.
The list of animal species, although variegated, presents no
zoogeographic novelties. Most of the species are characteristic of the
region in the last century, prior to the extermination of deer, chamois and
goat on the Skalistyi ridge. The find of suslik bones in the Mesolithic beds
is particularly interesting as related to the question of the age of the
populations of this species in the mountains of Caucasus. However, these
bones may have originated in the steppe plains of the Manych area and been
introduced into the layers from pellets of eagle owls and predators flying
south.
The Mesolithic site of Chokh in inner Dagestan was discovered
in 1955 by Kotovich 2 km north of the village of Chokh in the Gunib area.
194 The site is located under a steep limestone rock (Figure 86) on the right
bank of a small stream, which cuts deeply into a mountain ravine covered
187
(192)
FIGURE 84, Sosruko grotto in ВаКзап gorge
Photograph by S,N, Zamyatnin, 1956
1704 188
Бу an alpine meadow. The thickness of the Mesolithic layer in the excavated
areaexceeds 0.5m. The cultural layers contain flint tools in all stages of
finishing and finely-fractured bones. Among the flint implements are tools
with blunt backs, points, knifelike blades [or flakes] and various microliths.
(193) TABLE 41, Mammalian species and number of bones in Mesolithic beds of the Sosruko grotto in
Baksan gorge
Middle
Ages
Mesolithic beds
Early
I Age
Species
Domestic
о ео о а
LBS CHUM 5 5555550005500
Small cattle
ооо
Subtotal
Wild
WHgyes УИ 60505500
MARECS Ч oo o0080000000
MISS МЕ e565 co > oc
PaMMIErA раз 55600006
LEPWS SULOMEEGWS 45 as. ere
Спее В PwWFMASCMS 55 044% 1
Apodemus sylvaticus., oy
во Са бро =
MiEROKUS ШО оо оововоев =
SUS SCROWE доноров dine coc
Capreolus capreolus ... 1
CErnvyus ЭПБ sob >oo0 cn 6 ins
Rupicapra rupicapra . 1
CADW Clip Ch WCHSICa ие, oa
Artiodactyla (no closer
оО) оо бооюв оо J
БИОСа Gosctsane 50
ао оне оо со олени 50
The state of preservation of the fragmented bone material is similar
to that of fragments from beds of the third and second millennia B.C. in the
foothills of Dagestan. Identifications of the bone material in the 1955-1957
collections are given in Table 42 and Figure 87; most of the fragments are
unidentifiable. The rodents are represented by species which are
characteristic of the region in the present. They were introduced into the
site from the pellets of predatory birds. Amongthe ungulate bones, the most
interesting are those of a bison and a small ram, probably a successor to
the rams of the Lower Paleolithic Kudaro site.
189
195
FIGURE 85. Mammalian bones from the Mesolithic layers of Sosruko grotto
1 — jaw of Panthera pardus; 2 — дам (x 2) of Citellus pygmaeus; 3 — jaw (X 2) of
Ellobius talpinus; 4—jawof Cervus elaphus; 5 —astragalus of Capra caucasica
From the Koban burial grounds of the Bronze Age in Ossetia,
archaeologists longagocollected sculptures of local domestic animals.
These sculptures were made of bronze or, more rarely, of gold and silver.
Pendants, fibulae, metal plates, buckles and heads of staves were often made
by Koban craftsmen in the forms of heads of Caucasian goats, deer, elks,
wild and domestic rams, bulls, goats, horses and bears (Uvarova, 1900).
The past occurrence in the Caucasus of some of these species was
confirmed by our studies of Ossetian burials. The materials from those
sites indicate a late stage of existence of a rich assemblage of mammals
on the northern slope of the Greater Caucasus.
Dzuars of North Ossetia
Collections of skulls of domestic and wild ungulates from ritual sites
in the Caucasus have long been known. These collections were mentioned
by 17th century European travellers in Circassia (Jean de Luc, 1879).
Reference is made to them inthe notes of Guldenstaedt at the end of the 18th
century (Pallas, 1831), and in papers by Dinnik (1890a) and Satunin (1914).
The period of accumulation of skulls in the dzuars extends over the last
five centuries.
190
196
TABLE 42, Mammalian species and number of bones from the Mesolithic site at Chokh in Dagestan
Number
of bones
Number
of bones
Species Species
Domestic (? ) Capra sp. (cf. aegagrus or
Siva Со УИ ФО В) 46500506 8
Madd оо ооо Бюро босо а и, 94
Wild ааа oe closer
Па СЕМ) oid bo ob on Sao в 1,202
MIS UWE Боро обо ho 0 Bison bonasus (caucasicus?) 20
Small fragments of skeletal bones,
mainly Artiodactyla..........
HENS SWROPVEBOUS Ghooanoaeas
WMIGSOECHIOCSTOS AWWA DIST G 515.6 bo
Male Oly вере Ss 6 bo qb oo 60
Sis ЗСО о И th ON ОО о О И
Seong! CileyoiMmls’ еее
OWis go. (eh, вито Ил) .55064-
In 1947 and 1948 we studied 18 dzuars in the ravines of Urukh, Ardon,
Fiagdon and Gizel'don (Figure 88).
Skulls from dzuars located on the piedmont plains, e.g., the sacred
Khetag grove, were takenin the thirties to bone-calcination plants. Up to
30% of the dzuars in the mountain zone were also emptied of their bones
(Vereshchagin andSemenov-Tyan-Shanskii, 1948; Vereshchagin and Naniev,
1949).
The Adygeians, Digorians and Ossetians built their ritual sites in
picturesque groves onthe spurs of ridges, under rocky overhangs and incaves.
Special buildings were also constructed: churches, chapels or large
enclosed sheds made of stone or of large logs.
Dzuars were dedicated to different deities and each one belonged to a
particular village; the villagers assembled there for feasts on special annual
holidays of spring, summer or fall.
The custom of killing animals in the dzuars during feasts survived from
pagan times in Abkhazia. According to Abkhazian legend (Veidenbaum,
1879), a white bull used to emerge from the Oggin cave and was sacrificed.
In addition to ritualistic killing of domestic hoofed mammals, the
Digorians and Ossetians used to bring skulls of wild, hoofed animals to the
sites. This custom of bringing skulls into the dzuars was connected with
the belief in the god Avsati, protector of hunters and game, to whom the
skulls were dedicated. The occurrence of bison leg bones, with hoofs intact,
in the deep skull-bearing beds of the Digorized cave suggests that special
hunting expeditions set out before the holidays to bring an intact deer, elk
or bison into the dzuar. Skulls of wild, hoofed animals, boiled by hunters,
were also brought into the dzuars without relation to holidays. They were
either carefully laid in one common pile or on special shelves and supports,
or they were hung on cut branches of trees and placed against the walls of
the dzuar. It is difficult to estimate the age of the collections from the
number of wild animal skulls.
The state of preservation was not uniform among the skulls of wild
hoofed animals from different dzuars, because of differences intreatment
191
197 and preparation. No skulls had intact lower jaws; of the bison skulls, only
the hardest frontal and occipital bones with horn stems attached, were
preserved. This mode of preservation is identical to that of the skulls of
primitive bison from the shoals of the Don, the Volga and the Ural.
y V.G. Kotovich
FIGURE 87. Mammalian bones from Mesolithic beds of
the Chokh site
1-3 — First phalanx, metatarsal epiphysis and astragalus
of Ovis cf. gmelini
Only a few skulls were preserved in their entirety with basal part,
intermaxillaries and horns intact. The horns were fastened to the stems
with iron arrowheads inserted into holes prepared for this purpose. Horns
were also removed from their stems and used as wine goblets. Boiled
skulls of turs, chamois and roe deer were usually brought in after the basal
192
о Prokhladnaya
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о Shanaevo
о Beslan
‘=Tarskaya |
— — Fasnal i — :
A Styr-Digor eat г Aa т =
iz 9
Nogkau Dunta
э
а:
т %
= 75
и Gs
Oh cal WW
Mt,
HEELS
FIGURE 88, Map of some dzuars in North Ossetia
1 — remaining forests; 2 — ritual sites studied; 3 — limits of distribution of deer in the 19th century;
4 — area inhabited by relict bison and elk at the end of the 18th century; 5 — Recent glaciers
193
part had been removed with an oblique cut. The horns were removed,
together with skin and parts of frontal bone, from the fresh skulls of male
deer and elks killed in the fall. The brain was then removed through the
198 skull opening. The skulls were boiled, cleaned of flesh and small bones
and placed in the dzuar. All the skulls of female deer and elks are without
occipital bones which were excised to the dorsal mid-height. Most of the
deer skulls did not have nasal parts, as they separated from the skull in
the boiling process.
The most remarkable dzuars are Digorized (Olisai-Don), Lesgor I and
Lesgor II and Rekom.
FIGURE 89, Digorized cave in Jurassic slate cliff in Urukh gorge
Photograph by author, 1947
Digorized cave is located on the right bank of the Urukh gorge in
the southern cliff of the Skalistyi ridge. The cave was formed in the process
of the weathering of Jurassic limestones. Ledges, picturesquely covered
by pines, juniper and dog rose, occur on the giant cliff.
The cave is approximately 1,200 m above sea level. It lies within the
village limits of Zadelesk, located 1.5 km to the south.
The mouth of the cave is triangular in shape, approximately 12 m high
and 5.5 m at the base. The entrance is covered by a strong wall, 2.5 m high,
made of limestone fragments (Figures 89 and 90).
The feasting hall is approximately 7 m wide and 12 m long and contains
a primitive fireplace, narrow benches and tables made of split pine logs,
194
199
an iron cup for donations, two knotty-wood trunks placed against the wall,
carved wood fetishes hanging on the trunks and some utensils, consisting
of two large meat bowls, a wooden spoon and about fifteen horns of bison,
used as goblets. A pile of skulls and horns in the far left corner is heaped
between two fallen limestone blocks. A support, 2.7 m high, built of logs
and boards, stands in the forepart of the hall. Some 140-159 pairs of
antlers are piled on the support (Figure 134). The walls of the hall are
covered with resinous soot from the fireplace. The entrance to the narrow
right branch of the cave is blocked by an upright wooden log with a tin icon
of St. Nicholas. At the entrance, the height of the branching chamber is
about the height of a man; the height decreases and the chamber ends eight
meters from its entrance. The floor is covered by a meter-thick layer of
skulls and horns of artiodactyls (Figure 91).
The annual feasts in the Digorized cave took place in June during the
mowing season.
\y
\y (\ OY
KY \\" 0
Су Way
yp
FIGURE 90. Map of Digorized cave
1 — fallen blocks; 2 — fireplace; 3 — veremonial pottery; 4 — benches;
5 — support with deer antlers; 6 — stone wall; 7 —skulls of domestic
goats, roe deer; e — elk skulls; b — bison skulls
195
200
201
On our first visit to the cave in August 1947, three elk skulls and
15 bison skulls were found in the front rows of skulls on the floor of the
branch. These were probably the skulls noted by Guldenstaedt and Dinnik.
Dinnik mentioned three camel skulls which were not in the cave; he probably
misidentified the elk skulls. The pile of skulls in the left corner of the
feasting hall was covered by centuries-old dust layers, whereas the skulls
in the right branch of the cave had evidently been recently disturbed by
archaeologists or tourists. As the layer of skulls was being sorted, bent
or looped twigs (symbolic yoke), planed, half-burned sticks (shashlik spits),
iron arrowheads (Figure 92) and small silver coins were found.
The greatest number of bones found at Digorized were skulls of sheep,
and, in descending order, skulls of deer, large cattle, domestic goats,
bison, roe deer, chamois, elks, Caucasian turs, wild boars and buffalo.
The ratio was established by the value placed on the hunters' sacrifices
and by the availability of various game species in the hunting area around
the village of Zadelesk. The bison and elk skulls are the best-preserved
bone material because they remained on the floor of the dry parts of the
cave, between the entrance and the side branch. In many cases, ligaments
on the anterior and posterior sections were preserved because of the
absence of hide beetles. On the frontal bone of one of the bison skulls the
figure ''1833'' had been engraved with a knife. The elk skulls were in the
same state of preservation. Remains of tree twigs and bushes were well
preserved in the crevices of the animals' teeth. No elk horns were found
in the cave. The horns had been removed from the frontal and occipital
parts of one of the elk skulls inthe front row. In another skull, found in a far
corner, the horns had either fallen or been broken off the crown. It is
possible that the horns were stolen from the cave or retained by the hunter
as a particularly valuable trophy.
Deer antlers, found mixed with skulls in places protected from rain
and snow, retained their natural brown color. Traces of molting were
preserved on only a few antlers of deer which had been killed in August.
Most ofthe bones left on the support, however, became uniformly whitened
on the upper surface by rain, snow and sun, and blackened underneath
by soot. Skulls of chamois and roe deer occur mainly in the middle and
far section of the bone layer. Their state of preservation was much the
same as that of the bison and elk skulls.
The Dagestan goat is represented by four skulls with preserved horns;
their condition indicates that they were middle-aged. The three skulls of
wild boar found in the upper layers piled to the side can probably be
accounted for by the fact that the Digorians in later times abandoned their
earlier religious prejudices against this animal, and also by the fact that
the supply of more valuable animals dwindled. Fresh-boiled skulls and
limb bones, fractured by an ax, and ribs were only those of large cattle,
goats and sheep. It was clearly observed that large skulls of ancient,
mature bulls had been replaced in recent time by skulls ofsmall, immature
specimens, which had been broken longitudinally.
The skulls indicate that the large cattle of the region belonged to only
one variety during the entire period of accumulation of the bone bed. These
were large animals, with a concave area between the horns, which extended
forward, sideward and upward. The cattle belonged to the secondarily
enlarged European Bos brachyceros. Skulls of goats and sheep
196
(200)
FIGURE 91, Interior of Digorized cave
Photograph by author, 1947
represent relatively small varieties with
poorly developed horn stems and horns.
If it is assumed that, from the beginning
of the annual ritual, one bull and two to
three sheep were sacrificed each year,
it follows that the collection accumulated
over not less than 500 years.
The skulls in the right branch of the
cave were deposited over a thin layer of
limestone debris and old dust, underlain
by a solid plate. A reconnaissance trench
dug in the left corner of the feasting hall
down to the limestone floor (a depth of
60 cm) showed that there are no bones
in the layer of old dust.
Because skulls of bison and elks were
particularly valued by the Digorians as
ritualistic offerings to Avsati, it would
rhe aera seem likely that most of the skulls of
| р ст animals killed by the Zadelesk hunters
were deposited in the cave.
It is clear that the incidence of deer and,
particularly, of elk was very low even
as early as the 15th to 16th centuries. In
all probability, the period of accumulation
of the skulls could not have begun before the
Le MM Lh 127
FIGURE 92. Iron arrowheads from
Digorized cave
197
202
14th or 15th century and must have ended in the 18th or early 19th century.
The accumulation in the cave of deer skulls and horns, as indicated by their
state of preservation, ended only a little later than that of the skulls and
horns of elk and bison. However, the deer survived until the time
when the Ossetians obtained rifles. The Zadelesk hunters killed on an
average up to ten deer each year.
The Digorized bone material is particularly interesting for analysis of
the age and sex groups of the game, as well as the seasonal hunting yields.
These data are given in Table 43. *
TABLE 43, Sex and age groups of wild ungulates from Digorized cave
Females
Species
immature | mature total
От ieee»
Capra cylindricornis ....
Rupicapra rupicapra
G ANU GIAIS CAs. 87a seneakt ee age ey = 10
Bison bonasus caucasicus 30
Capreolus capreolus..... 34
Cervus elaphus, шага... 439
Areas асе Cail cas EWS ae
The main game animal, deer, was hunted mostly in the fall, during
the rut.
Dzuar Lesgor I (lower) was built on a small ledge of the same
Skalistyi cuesta and at the same altitude as the Digorized cave, but on the
left wall of the Urukh gorge. It belonged to the village of Lesgor, which
was.abandoned at the beginning of this century. The skulls were arranged
in two layers behind a stone fence on an open wall. The lower layer rests
directly on the ground, the upper on a big pine log inserted into a crevice
in the rock, and on a "'hanger'' made of a knotty-pine trunk. The upper pile
of skulls is colored by the smoke of the ritual fire, which was regularly lit
under the rock. Most of the skulls and antlers of deer in the lower layer
decayed because of moisture. The following kinds of skulls (given in
descending order of occurrence) were found: sheep, cow, goat, deer, east
Caucasian goat, roe deer, bison and elk. Among them was a completely
'fresh'' skull of a young bison and a completely smoked skull of a she-bison
with right horn intact.
Lesgor II (upper) is located 250 m higher on the cliff on the side of
а very steep path in a picturesque crevice. The collection of skulls is
under a small rock overhang and is poorly protected from weather. Deer
antlers rest on a horizontal, elevated support made of pine logs. Some of
the skulls on the ground have ''grown"' into the soil and become covered
with sod. Skulls of wild, hoofed mammals are more numerous than at
Lesgor I, probably because the dzuar is located closer to the hunting grounds
* Only the skulls whose age and sex were identified with certainty were counted.
198
203
and farther away from the village. The descending order of occurrence
is as follows: sheep, east Caucasian goat, deer, cow, goat, bison,
chamois, elk and roe deer.
As at Lesgor I, the horns are missing on most of the goat skulls; they
were broken off and probably taken to the village (Figure 93).
This group of the Urukh dzuars is very important in the study of the
past ranges of deer and elkand their gradual shrinkage. Since it was
difficult to carry the skulls and horns by mountain paths to the dzuars,
hunters chose the most venerated dzuars and those which were closest to
their hunting grounds. From the location of the ancient mountain paths
through the passes of the Skalistyi ridge, which connected the villages
Zadelesk and Lesgor with the piedmont plain, and from local legend, it can
be established that most of the skulls of deer, elk and bison originated
below the dzuars in the forests of the Chernye gorge and on the sloping
Terek plain, particularly in the vicinity of the village of Akhsarisar. Even
inthe 'twenties, deer still appeared during the mating season in the upper
Urukh gorge, near the villages of Styr-Digor, Nogkau, Kamunt and Dunta.
The last of the bison and elk, however, mainly inhabited the foothill forests
of Ossetia toward the end of the 18th century. This is confirmed by the
absence of skulls of these forms in the upper dzuars. In the longitudinal
valley proper, between the Skalistyi and Bokovoi ridges, deer, elk and
bison were probably either exterminated or driven out before the time of
the Digorians by dense settlement and destruction of forests by earlier
inhabitants. There are reasons to believe that elk and bison still frequented
the forests of Ossetia at the time of Guldenstaedt's travels, although only
sporadically.
FIGURE 93. Skulls of east Caucasian goats in Lesgor II dzuar
Photograph by author, 1947
199
Rekom Dzuar is the best known dzuar in the literature. It is
located in the gorge of the Tseya (a tributary of the Ardon) in the pine-
forest zone of the Bokovoi ridge and is greatly venerated by the Ossetians
(Vyazigin, 1929). The dzuar-belongs to the village of Tsei, but is visited
by villagers from other places as well.
The dzuar is a massive structure dating from the end of the 17thcentury.
It is constructed of pine and yew logs with one door and no windows and is
situated in a pleasant meadow surrounded by old pine forest and rocks,
with an outlook to snow-covered peaks (Figure 94). The site was inhabited
as early as the Bronze Age.
Those skulls which remain after they were plundered by tourists for
souvenirs rest on specially constructed wall ledges and shelving of raised
earth. The material consists of skulls and horns of Caucasian goats,
domestic sheep, cows, goats and deer. (Satunin (1914b) reported bison
skulls from the Rekom dzuar.) Most of the remains of Caucasian goat
consist of horns and soft horn stems weathered and split by moisture and
sun. Six fresh-boiled skulls were also found, brought probably from the
margins of the nearest glaciers during the last 20-30 years. Chamois still
live in the rocks above the dzuar. Because of the deep winter snow deer
and roe deer probably did not inhabit the region even in the past. Most of
the deer skulls and horns of Rekom were brought there from the lower areas
of the Ardon basin.
204
FIGURE 94, Rekom dzuar
Photograph by author, 1947
200
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206
207
The horns and skulls of Caucasian goats belonged to mature or even aged
individuals of 6-17 years; the most common age is 10-11 years (Figures
95-96).
From the 18 dzuars we collected up to 10,000 items: skulls, horns and
fragments of skulls and horns of 8,815 individuals. They belong to 11 species
of hoofed mammals: domestic species of bull, buffalo, sheep and goat; wild
species of boar, bison, east Caucasian goat, chamois, elk, deer and roe
deer (Table 44).
Our studies of the skulls in the Ossetian dzuars compared with data
drawn from the literature and with other verified data lead to the following
conclusions:
1. One and a half centuries ago the Terek-Sunzha inclined plain and the
canyons on the northern slope of the Greater Caucasus were inhabited by
eight species of artiodactyls: boar, bison, east Caucasian and west
Caucasian goats, chamois, red deer, elk and roe deer.
2. Bison and elk were on the verge of extinction by the end of the 18th
century, at which time deer were still abundant.
3. The final extermination of deer in that section of the Greater Caucasus
under study occurred in the second decade of the 20th century.
4. The number of goats, chamois, roe deer and boars greatly decreased
during the last century and the beginning of the present one.
5. It was chiefly human activity that brought about the extinction of some
species and the population decrease of others, directly through unrestricted
use of firearms and indirectly through the cutting down of forests on the
inclined piedmont plain.
Ritual sites similar to those described earlier, containing bone
material mostly consisting of skulls and horns of east Caucasian and wild
goats and deer, are also known from Khevsuretia, Tushetia and Dagestan
(Dinnik, 1914a; Makalatiya, 1935; Maruashvili, 1955).
Bones of small mammals in pellets of eagle owls and little owls have
been collected by the author at two localities in the inner valleys. These
bones occurred at the bottom of small caverns in limestone rock near the
village of Dzivgis in the Fiagdon gorge. They are not more than 150-200
years old, completely white and in a good state of preservation.
The present landscape of this area is composed of turfy dry slopes
covered with tragacanth, astragali and rocks. Vegetable gardens, hay
meadows and pastures are located near the river. The height above sea level
is approximately 1,400 m. The list of the fauna is given in Table 45.
Approximately 40 fragments of lizards and toads were collected at the
same locality.
All the species mentioned live in the present in the dry part of the
Fiagdon valley. However, it should be mentioned that the occurrence of
Norway rats, domestic mice and, possibly, migratory hamsters are
related to the deforestation of the valley.
Bones of other species were found in the pellets of little owls in the
ruins of a medieval fortress on the upper reaches of the Doniserdon, the
right tributary of the Urukh.
The upper parts of the longitudinal valley are covered with mesophytic
meadows and birch groves. The altitude is nearly 1,600 m above sea level.
The fossil species correspond to the fauna of the Recent landscape (Table 46).
202
A comparison of Tables 45 and 46 which takes into account the evolution
of landscapes in the inner valleys indicates that the composition of the mice
fauna and their distribution in the dry inner valleys changed considerably
over the last few centuries.
10
Frequency
GEO) OU bon N70 Vara 6 90а 95
Horn length (cm)
итал fou 17 в
Age in years
FIGURE 95, Relationships between age groups and size of horns of east Caucasian
goats in Rekom dzuar (length of horns measured along the maximum curvature )
(206) TABLE 45. Species and number of bones of mammals from pellets of eagle owls
and little owls near Dzivgis village
Number
of individuals
Number
Species
В of bones
Vulpes vulpes
eC
Mausiiediayenpiwy а ме swam 2 jah 1
IL SVMS Sw OPAEWS 45.044 ole bio or 1
(Gwi oe ew ws ата TEEN WS ARYA a 22
М ево EWES 355.46 5 also #
MWS ИОВ 5 500400000000 3
Apodemus sylvaticus ....... 6
RALEOS MOVES И sondage o eo 7
203
The increasing aridity in the valleys, caused by cutting the groves оп the
slopes and grazing of cattle, resulted in the replacement of the mesophilous
ecological assemblage of rodents (Microtus arvalis, M. majori,
Chionomys gud) by a xerophilous assemblage (Cricetulus
migratorius, Mus musculus hortulanus, Lepus europaeus)
and by the introduction of synanthropic forms (Rattus norvegicus,
Mus musculus musculus).
FIGURE 96, Horn covers of goats and deer antlers along the
walls of Rekom cave
There are no data on the recent extinction of small mammals in the
areas under study. Also no remains were found of those ancient species
of little suslik (Citellus pygmaeus musicus) and Radde's hamster
(Mesocricetus auratus raddei) which are characteristic of the
inner valleys of Kabarda in the west and Dagestan in the east.
208 Western Transcaucasia
In the Holocene human settlements clustered around the terraces of the
Black Sea coast, mountain river gorges and the shores of water bodies in
swampy Colchis.
204
TABLE 46, Species and number of mammal bones in pellets of little owls from the
upper Doniserdon
р Number Number of
Species Tene
of bones individuals
Зотех DREUIO edie me ое, 5 1
Nal DUG Call aha EIT о songs опа 16 1
МОЛОТ BWC Gos sb dodo 600 cc 37 6
МООИ © Geb obboo0ome 0000000 727 80
Including the number identified with
certainty:
Mi main via TiS, вона = (77)
Манон ое eae ae ees god (3)
MWS ЗО И о 666004 dd oo Ho 6
TABLE 47. Species and number of bones of mammals from post-Paleolithic beds of the Akhshtyrskaya cave
Number of Number Number of
Species individuals ee of bones individuals
Domestic Wild
О. Бе Ш 222809 oo 54050 2
О G13 Wee odo оо Camis Пиры еее 1
Ри еее MIBGIES ее 5455665 1
WARES CAMS 556000 Lepus europaeus... 1
COLIN об Dace eo oot RAETWS РА Е ообооб 1
Е LSS Aap cla ow Зичаенога Ба Л: ме 4
Esrviis Claplims соо - 1
Повар. wae. Capreolus capreolus 5
Subtotal
The fauna of the Neolithic and younger beds of the Akhshtyrskaya
cave near Adler are of particular interest. These beds overlie the
Paleolithic, but it is difficult to subdivide the material from the upper
layers of the cave (Zamyatnin collections) into the various post-Paleolithic
209 cultures. The bones are light in weight, mostly light yellow in color with
some dark stains. Some of the fragments have partially lost organic matter;
in general, theyresemble Upper Paleolithic bones. The identifiable material
consists mostly of epiphyses. The species identified and their number are
given in Table 47* (Figure 97).
* The material from the Neolithic and Iron Age layers was combined, because the state of preservation
of the bones definitely indicates that they were mixed in the layers and during the excavations,
205
210
\ \ ШИ /
i i |
!
—
/, Hy! ] ‚ |
Yi Min \\ \
Se
| Wy. :
FIGURE 97, Mammalian bones from post-Paleolithic layers of the Akhshtyrskaya cave
1 — metacarpal of Ursus arctos; 2—radius of Meles meles; 3—femur of Lepus
europaeus; 4—femur of Rattus rattus; 5—metatarsal of Capreolus capreolus;
6 — metacarpalof Capra caucasica
The composition of food remains at this site is more variegated in wild
fauna than at other post-Paleolithic localities in the Caucasus, since the
cave was probably used by hunters and shepherds.
The absence at the site of mountain ungulates (cave bear, elk, giant
deer and bison) which were probably driven farther into the mountains is
quite apparent, as is the appearance of black rat in the Iron Age beds.
In swampy Colchis the oldest cultural beds containing animal remains
are known from the excavations of an Eneolithic site. The settlement was
on a mound, at present submerged in a swamp, near the village of
Ochamchiri, on the left bank of the Dzhikumur creek (Solov'ev, 1939).
In the bone collections Gromov has identified a small cow, domestic pig,
goats and sheep. Wild species include deer, roe deer and fish, particularly
flounder. It is characteristic that horse is not found among the remains.
One of the Early Bronze Age settlements whichis located near Anaklia,
close to the mouth of the Ingur River, has been excavated by archaeologists
of the Zugdinskii Museum. The cultural layer is overlain by Recent silty
alluvium. According to Kuftin, the very primitive black ceramics place
this bed in the second millennium B.C. The bones from this site are
shiny, chestnut in color and slightly fractured. The latter characteristic
indicates that the ancient settlers of the coast had an adequate supply of
206
animals for food. The species identified (Table 48) show that primitive
animal husbandry and more developed hunting were carried on by the ancient
settlers.
TABLE 48. Species and number of bones of mammals from the Bronze Age settlement near Anaklia
Species Number of bones Number of individuals
Domestic
Ро ИЕ a AUR a te 103 17
LEWES: CAME. оо оо ооо 0 oo dela обла в Ва 258 17
CON WEAR и Olg).o eG) cl oeb) А 47 5
ОА bao 009 00 ple 408 39
Wild
Grains) “AMEE SUS eee etal ее 1 1
ОО: S25 coco eoaocaonos odbc te 3
Sis: эст Offa И о 15 3
Сетунь GlApINNS ooboevobsoogdueo0 gan 4 1
(EelpoymGOUMS CAPFEOQMSS cacao oobacu neo 36 4
sub ee ee СИ РТ," 2
WOW oo soababoanponoadaos 468 51
Individual bones of fish and birds are also known from this locality.
At present deer do not inhabit the area near the site, but jackals are
abundant, and boars and roe deer occur occasionally.
The absence of horses and the Small number of dolphin bones, as
compared with the Gelendzhik locality, is characteristic of the Anaklia site.
Rich bone material, which has probably been lost, was obtained from
excavations of the numerous mounds and settlements of ancient Greek and
later times on the Colchis plains, along the midde and lower sections of
the Rion. The author has identified remains of a domestic goat, wild boar,
deer and roe deer, found at the Naokhvamy site, from photographs by the
archaeologist Kuftin.
A number of fragmentary antlers of red deer, obtained in the peat areas
of Colchis during the construction of drainage canals, were deposited in
the museum of the town of Poti.
Avery important deposit of food remains was exposed in the excavations
of Holocene (Neolithic and latest) beds at the Sagvardzhile site, on the
eastern margin of the Colchis plain (Kiladze, 1953).
The collections from these beds which have been studied by the author
contain the following wild species in order of decreasing abundance: *
Sus scrofa Ursus arctos
Cervus elaphus Equus caballus
Capreolus capreolus Felis silvestris
Bison bonasus Panthera pardus
Capra caucasica Castor fiber
Rupicapra rupicapra
* Judging by the state of preservation, this material, which is under study by Burchak-Abramovich,
undoubtedly also includes Paleolithic bones,
207
The occurrence of the beaver bones confirms that it was in Colchis that
this species survived the Pleistocene in the Caucasus.
Of the many western Transcaucasian monuments of ancient architecture
and sculpture ornamented with animal design, the Bagrat temple (1008, A.D.)
in Kutaisi is particularly noteworthy. A scene showing cheetahs and
panthers attacking a horse or a kulan and a goat is depicted on the heads of
the temple columns. Taking into account the broad base of medieval
Georgian culture it is obviously difficult to maintain on the basis of these
ornaments that kulans and wild horses existed in Imeretia in the 10th
century (Figure 98).
FIGURE 98, Scene on the head of Bagrat temple column — panthers attacking a goat
Photograph by author, 1956
The latest developmental stages of the ecological assemblages of small
mammals in Imeretia can be traced through bones brought by eagle owls
into small caverns in the walls of the canyons which cut through the
Imeretian Plateau. In 1952 we collected material (identified in Table 49)
from close by the Sakazhia cave inthe Tskhali-Tsiteli gorge which
shows the interesting, simultaneous occurrence of jackal, hare, Norway
rat and fat dormouse, species not known from the Paleolithic beds of
Imeretian caves.
As opposed to eastern Caucasia, no collections of game animal skulls
have been preserved from the mountainous section of western Transcaucasia
in the Ingur, Rion and Tskhenis-Tskhali gorges which were inhabited by the
Svans.
212 Radde, it is true, wrote in 1866 that in 1864 he had observed ап
accumulation of horns of Caucasian goats, piled by hunters near the chapel
of the village of Chibiana (Ushkul'), inthe upper reaches of the Ingur.
He estimated the number cof horns at ''several thousands." In 1948 I could
208
find only 7 pairs of horns and horn stems of goats (6 of Capra
caucasica апа 1 of C.cylindricornis) inthe altar of the chapel in
the village of Zhabeshi and in the altar of the Lamardiya monastery above
Ushkul'. No confirmation ofthe past existence of ''thousands of goat horns''
could be obtained from old residents or from the local authorities. But it
seemed that the hunters of this village used to store horns and blade bones
of Caucasian goats and chamois under the roofs of their observation towers.
By 1948 the collections were mostly abandoned or given away and no new
material was introduced due to the extermination of the animals.
TABLE 49, Species and number of Holocene bones of mammals from pellets of
eagle owls in the Tskhali-Tsiteli gorge
Number of
individuals
ss a Number of
cies
р bones
EriMmACCUS SUTODAGCWS 5452000 4
Camis ЕСО Fo gioco 6 aieoetono 1 1
[ею CURODASCWS оооовобеов 2 1
ооо о бора Chal oMaNE NS 12 3
2
RECESS МОУ МВ Gooeanos
eet ee we we
It was the practice of the Svans, as opposed to that of the Ossetians,
to cut the goat horns off with only a small piece of frontal bone. Evennow
goat horns and bear paws can be seen attached to the porches of houses
and embedded in the stone walls of houses made of slate.
Thus, the studies of Holocene bones in western Trancaucasia indicate
an abundance in the Neolithic and later epochs of larger game animals:
boar, deer, roe deer, bison andbear, as well as the European beaver found
in Colchis. The appearance of new species, unknown in the Pleistocene
(Paleolithic) beds (jackal, Norway rat and black rat) is highly
characteristic. The composition of the domestic fauna and the nature of
their deposition indicate that, ingeneral, the landscape of Colchis andthe Black
Sea coast was very Similar to the landscape of the beginning of the present
century.
сео аа саж е ата
Reconnaissance excavations were carried out in 1952 by Lyubin above
the overhangs of the natural landmarks of Saroty-Or and Sharshiat-Kadzakh
near Staliniri. We have identified in Lyubin's collections bone fragments
of small domestic pigs, bulls, goats and sheep, and teeth of European brown
bears and foxes. Rich collections were also obtained in 1955-1957 from the
Holocene beds of the Kudaro caves on the upper Rion.
In the valleys of the Kura and the Aragva rich bone material was
excavated by archaeologists of the Caucasian Museum from multilayered
209
213 cemeteries andtowns at Samtavro and Armazi, near Mtskheti.* А сотраг150п
between photographs of this material in the archives and exhibit material
of the Upper Bronze Age (9th-8th centuries B.C.) show that these were
bones of horses, cows, sheep, goats andboars, and amulets made of canines of
European brown bear and incisors of beavers. Skeletonsofmartens (Martes
foina) and steppe voles (Microtus socialis), which died at a later
period, have recently been discovered. Objects made of red deer antlers
are very common, e.g., the so-called ''spear rectifiers. '' Elegant bronze
plates and trinkets of this period from Samtavro and Armazi often depicted
dogs, foxes, lions, panthers, hares, deer, goitered gazelle, rams,
Caucasian goats and Bos primigenius.
Taking into account the paleontological data for Transcaucasia as a whole,
these artistic representations confirm the existence of some large game
animals in the Caucasus during the first millennium B.C. These animals
became extinct only in the Middle Ages.
Bastern D2rans caucaisi a
On the Kura Plain and in the foothills of the Greater Caucasus, Karabakh
and Talysh, Holocene deposits of bones occur in diluvium, alluvium, caves,
asphalt crusts and cultural layers of ancient settlements.
There is a record of a horn of a red deer from the diluvium of a mountain
slope near the village of Alty-Agach, near the eastern margin of the
Bolshoi Range (Deminskii, 1901). Many occurrences are concentrated on
the Apsheron Peninsula.
Kovalevskii found in 1941 a skeleton of a striped hyena (Hyaena
hyaena) at a depth of 4 m in Holocene loess loams in the lower section
of the Sumgait rivulet, north of Baku. The skeleton was buried undera
cultural layer containing medieval pottery and under a bone-bearing bed of
an older settlement, overlain bythe sediments of the Recent Caspian. From
the stratigraphic position of the skeleton and its state of preservation
it can be dated as Lower Holocene.
A skull of a very old striped hyena, bones of a horse and of a Caspian
seal (Phoca caspica) were found by Kovalevskii in the Bailovskaya
cave onthe southern outskirts of Baku (Bogachev, 1938b).
We found two leg bones of goitered gazelle (Gazella subgutturosa)
in blocks of asphalt-impregnated surface loams overlying the Middle
Quaternary beds at Binagady. The bones are fresher and less impregnated
with oil than the Pleistocene bones. In the asphalt crusts of Mount Kir-
Kishlag we discovered bones of hare and wolf. Skeletons of small rodents,
jerboas (Allactaga elater), hamsters (Cricetulus migratorius)
and voles (Microtus socialis), which died in asphalt, also occur in
this locality and at Mount Zigil'-Pirya (Vereshchagin, 1951b).
The numerous Holocene settlements and burials of Azerbaidzhan (Passek
and Latynin, 1926) are responsible for the rich bone collections from the
cultural layers.
Gummel!' (1948), who excavated the Bronze Age mounds on the sloping
plain west of Khanlar (formerly Helenendorf), mentions numerous bones of
* This material remained unidentified and was subsequently lost.
210
215
Bos, sheep, goat and domestic pig. The number of horse bones was
negligible.
Excavations of the settlements and the burial near ancient
Mingechaur оп the right bank of the Kura, near a hydroelectric
power plant, revealed cultural layers dating from the second millennium
B.C. to the early Middle Ages. The very rich bone material from this
locality has not yet been completely identified. Our preliminary study of
these materials indicated the presence of numerous bones of horses, asses,
cows, sheep, goats and, more rarely, domestic pigs. The wild animals
are represented by red deer (Cervus elaphus), goitered gazelle
(Gazella subgutturosa) and boar (Sus scrofa), all of which were
until recently quite common in the region.
We found a particularly interesting burial near Mingechaur with bones
of black rat (Rattus rattus) andhorse, ass, large cattle and
boar buried in jugs. Thistype of burial, which included sacrificed domestic
goats and grain, is usually dated as middle of the last millennium B.C. In
view of the fact that the burial field has been covered for many centuries
by a semidesert saltwort vegetation, the rats could only have inhabited the
field during a time when it was settled and irrigated or at the time when
the cult of mass burial flourished. Their occurrence verifies assumptions
of the early appearance of black rat in Transcaucasia (Vereshchagin, 1949c).
TABLE 50, Species and number of mammal bones from ancient settlement near Sumgait
Species Number of bones Number of individuals
Domestic
FVOTS CMe, cere ts и Tans Elva eRe 6 1
ато еса ее 11 3
SHEED YU ен пани 12 3
SUID ALO aM age ие м 29 7
Wild
LEPUS МОИ о Бове boo boods bona
GaxvZeila SMD GUULWROSE oh оба Вбр в ails
2
3
а.
oO} NO] =
A very old settlement was discovered by Kovalevskii in 1941 in Holocene
loams near the mouth of the Sumgait rivulet. Food remains were
found at a depth of 3 m under the cultural layer containing medieval pottery;
they were well preserved under the very dry burial conditions. The species
found are given in Table 50.
At the same locality ten fragments of bones of mute swan (Cygnus
olor L.) and goose (Anser anser L.: three specimens) were found.
According to Kovalevskii, the bone-bearing bed is overlain by sediments
of the present-day Caspian and therefore antedates the last Holocene
transgression of the Caspian, i.e., early first millennium B.C.
211
216
Large collections of bones were made in 1953-1954 by archaeologists of
the Azerbaidzhan Academy at the excavations of the fortress Uren'-Kala
(Bailakan) in the center of the Mil'skaya steppe. We observed the layers
from the Middle Ages at this locality to be characterized by sheep, large
cattle, goat andhorse, witha small admixture of goitered gazelle (Gazella
subgutturosa).
TABLE 51, Species and number of mammal bones from the medieval settlement
on the Baku Fortress hill
Number of
Species Number of bones
P i. individuals
Domestic
a) ое: |) else) мы. м ale (a) Plone) 160) 81 Ге»
о ед te) iw, ое Ole) ual Pe) (@) \¢. (8) аа
Oe Че 1s) 6) fe) fe) we 6 о @ ere "Че
ча 5 folle обеде fel ere ру. 0s 0, НА e ©
dic’ s ee «Le bw > eke io iota $ и
fede. ви в м & 0) 6) See Rees Ba oi te
Se oS, м опер Oe (6 оюеае
ОР OO DOs, ome ONCE Om) Cer rar
7 ОМ Cree: СО oc
"ОЕ ЕТО pun ding Э We ons Eel ай
NAGY SSA Hh SOMOS Bis еее &
вания Wem Onis! а се сь
Gazella‘subgutturosa ......,
Sos бя виа sue и
Ble) 640, we 2 e100; ae
Bone material from beds of the Middle Ages was also obtained by
Leviatov in excavations of the Shirvan-Shah Palace courtyard in Baku.
The excavations revealed that small settlements successively occupied
the hill of the contemporary Baku Fortress. The beds which have been
studied are dated by the pottery as 9-13th century A.D. This is the time
of the end of the Arabic caliphate, the age of Nisami and the beginning of
the Mongolian invasions into eastern Transcaucasia. Part of the bone
material buried at the site probably resulted from epidemic death among
the cattle, since complete scapula and forelimb bones are common.
However, most of the bone fragments are food remains which were well-
preserved in the dry ground.
Bones of domestic animals — sheep, cows, horses and asses —
characteristically predominate. Bones of wild species — goitered gazelle,
kulan, fox and seal — are not numerous (Table 51).
212
211
FIGURE 99. Ruins of Chukhur-Kabala fortress
Photograph by author, 1951
From the same beds the following birds have been identified: Cygnus
Spey Аизее (sp., 5 Бава. © серва» № domestic fowl,
totalling 21 fragments of 10 individuals. In addition bone identification was
made of the following fishes: Acipenseridae (25 fragments of 4 individuals)
and Cyprinidae (58 fragments of 8 individuals).
Most of the goitered gazelle bones occur in the 10--11th-century beds.
The metatarsus of a kulan was found in the 13th-century bed.
The findings in ancient Baku confirm an abundance of goitered gazelle
and kulans in eastern Transcaucasia. The hunting of seals and aquatic
birds, andfishingin the Baku bay have also been established.
Bones and complete skulls of rats (Rattus norvegicus) are common
in recent layers, dated 15-16th century A.D. However, the contemporaneity
of the bones and the beds is not established, since rats may often penetrate
the deep layers by descending through crevices and holes in the ground.
We studied bone material from the beds of the ancient Chukhur -
Kabala fortress (Figure 99), located on the northern margins of the
Kura Plain in the Alazan-Agrichai valley and destroyed by the Mongolians
in the 14th century (Krymskii, 1934). The ruins of the fortress and the
towns are located between two left tributaries of the Turyan-Chai in thick
broadleaf forest. During the period of the maximum development of these
towns the trees were probably felled, since fragments of square bricks
and pottery occur in the vicinity, particularly in the beds of the Kara-Su
springs which flow through the alder and wing nut thickets.
213
218
Bones of wild and domestic animals, mixed with pottery, bricks, ash
and earth, fill deep wells in the fortress, which are at present being active.y
eroded from the east by the Kara-Chai rivulet. The identified bones of
domestic and wild animals which we collected in the talus of the cliff and
in the rivulet bed are given in Table 52.
TABLE 52, Species and number of mammal bones from the Chukhur-Kabala
fortress beds (Middle Ages)
Number of
individuals
Species Number of bones
Domestic
Large cattle ae ines yeild ate) ere elles
В Зе Ао ое Эва
arn No
Sin bitoit а ous ila
Wild
Glervus ета ох
Capreolus сартео 1 so а =
eee eee eee
Special excavations would undoubtedly increase the number of wild
species specimens. In 1946-1947 we collected bones of small animals on
the northern margins of the Kura Plain and in Apsheron from owl pellets.
The material was collectedbetween Aldzhigan-Chai and Geok-Chai
in short-livedcaves inthe cliffs of the Tertiary hills which separate the Alazan-
Agrichai valley from the Kura Plain. Study of the fossil material showed
that it was identical with the modern assemblage of insectivores and rodents
known from samples obtained with traps (Table 53).
We obtained more complete materials in a study of past and present
distribution of small mammals on the Apsheron Peninsula, which enabled
us to compare the composition of the present and the Pleistocene faunas.
A narrow crevice (Figure 100) which had been occupied for several
centuries by little owls was discovered by Burchak-Abramovich in 1944 in
the limestone rock east of Cape Bil'gya onthenorthern coast of the
Apsheron. Bones of insectivores, rodents, birds, reptiles and amphibians,
and chitin of beetles from decayed pellets formed a layer 10-12 cm thick
in the cavern. The bones in the upper layer were brown in color; those
in the lower layers were gray-brown. Gromov, in 1944, andI, in 1945,
collected up to 5,500 bones of mammals in the crevices.There were fewer
bones of birds, lizards and toads.
214
TABLE 53. Species and number of bones of small mammals in owl pellets from Geok-Chai
Number of bones
Species Number of individuals
Hemiechinus auritus ....... : 1
Соса пов о 5 466 bol uk 7
оо ИЕ Иа 6.5 6 S60 5 o Bip e 1
Cricetulus migratorius ...... 2
MACROS БО ЧЕ ИО поло se deo cc 19
MTS И В So WSs. koe нь 6
Арочетни 5 у маи що 5b o aos
MSTIOMES SRyWlarOUlrws 5569400 4
ee
FIGURE 100, Resting place of little owls in the rocks of Cape
Bil'gya
As old and plentiful as the accumulations found in owl prey are, they
contain an essentially modern faunal assemblage, very similar to that which
occurs at present on the northern Apsheron and is known from extensive
samplings obtained with traps (Table 54).
In both the owl-pellet accumulation and the trap samplings, there is a
disproportionate representation of certain rodent species, which reflects
215
in the one case the selectivity of the owls and in the other the specific
instructions under which the personnel of the anti-plague station operated.
There is no doubt that the ecological assemblage of smaller mammals
of the Apsheron remained relatively stable over the last part of the
Holocene. The study of the distribution of Recent mammals showed that
the European hedgehog, forest dormouse, field mouse and steppe vole are
relict species which have survived in a few sections of the peninsula from
a more humid climatic stage, as have some other vertebrates, in particular
Rana ridibunda WU. and the pond turtle Emys, orbreularmis L.
(Vereshchagin, 1949c) (Figure 101).
TABLE 54, Species and number of smaller mammals on Apsheron
Caught by owls in the
last three to four
centuries
Species Trapped in 1939-1940
ETinace us europaeus чо. .. ею Rare
Неттесн и AUT RAMUS ое сес Common
Crocidura russula.7. о, ее Common
Pachyura еттизса ..,.. а emer cee Not recorded
Юуто уз Whi Gost pac So sooo oy ooo Rare
Cricetulus, migra commie ene mack 73
Microtus socialis Я а 115
Meriones ery throupiisee ое о нь 2,123
А11астаса elaiter о Wwe и 8
А. williamsi ... .. ое tie 50
Mus musculus .. . ВЕ caer mea 81
A pode mws sy мати, о kao deel tale
Rattus norvegicus
* The number of individuals is estimated from the number of lower jaws.
The graphic art of nomadic tribes provides indirect evidence of the
occurrence of large mammals inthe historical epoch ineastern Transcaucasia.
A number of ancient campsites of nomads occur under the cliffs of
Apsheron limestone in Kabristan, southwest of Baku. Ancient craftsmen
have left line drawings of lions, red deer, wild and domestic goats,
domestic bulls and horses on the picturesque rocks and overhangs of Mount
Kyzyl-Kum, Beyuk-Dag and Dzhangir-Dag, which were probably also
religious sites (Vereshchagin and Burchak-Abramovich, 1948). Some of
the drawings are probably Neolithic, some Bronze Age and some belong
to the present era (Figure 102).
The studies of Holocene burials in eastern Transcaucasia may be
Summarized as follows. In the postglacial epoch the following species
appeared in eastern Transcaucasia: wild forms — striped hyena and
possibly lion; rodents — black and Norway rats; hoofed mammals — kulan
and goitered gazelle. A number of large mammals, characteristic of the
Middle and Upper Pleistocene, disappeared from the area probably at a
somewhat earlier stage.
216
49°00' 49°30' 50°00"
PRESENT DISTRIBUTION OF SOME
RELICT SPECIES ON THE
Ly APSHERON PENINSULA
Wy
Г =
Саре Bil’gya
М la
“2 rivulet У
=
Mt. Keklik- Dag pany,
"ify == М“ ое ae
#7 Рига <> Магоеп Г.
РА, Ел = Mt. KergeaJ
“Mt. Osman-Dag и” Rana ridibunda
aoe 7
` Be and pond turtle
1
: | polycarpos (1)
European hedgehog @ Juniper J.polycarp
(i bbionge (2)
малыш Rock areas with brush
Steppe vole on
Apsheron
eDuvannyi Г.
_ Sulla I, 40°
FIGURE 101
(220)
ой
Рае =
ры.
FIGURE 102, Drawing of goats and other animals on rocks of Dzhingir-Dag
217
221
Until recently some ungulates (е.5., deer and goitered gazelle)
were much more abundant; the deer inhabited the tugai forests of the middle
reaches of the Kura River.
The occurrence of relict species (European hedgehog, fat dormouse,
field mouse, Rana ridibunda and pond turtle) on the Apsheron
indicates that the climatic regime remained relatively stable during the
historical epoch. Their distribution also reflects changes in the landscape
caused by man.
Lesser Caucasus Upland
Postglacial mammals of the Armenian Plateau occur in diluvial
sediments, tuffaceous conglomerates, lake sediments, caves, cultural
layers of settlements and burial grounds.
Localities in diluvium on the shores of Lake Sevan.
As early as 1926 Shelkovnikov brought to the ZIN rounded fragments of a
bison skull (Bison bonasus subsp.), collected from postglacial,
tuffaceous conglomerates at the Zanga source in Lake Sevan (Gromova,
1935c). The state of preservation of this material shows that it may be
Bronze Age.
A large deer skullwithexcellently preserved antlers was collected from
the diluvium of a slope near the village of Kabakhlu in the Azizbekov area
in Armenia (Dal', 1947a).
A fragment of the skull of a suslik (Citellus cf. xanthoprymnus)
was collected from the alluvial loams on the right bank of the Zanga,
6.5-7.0 m below the surface, in the vicinity of Arzni. The loams overlie
a lava flow, presumably of the third to second millennium B.C. The time
of the suslik is considered to be somewhat more recent (Dal', 1949a). The
present eastern boundary of the suslik range is 40 km to the west.
Many animal bones were deposited in the littoral zone of Lake Sevan, some
from ‘animals which fell through the ice and drowned, some brought by the
mountain streams. A large proportion of the bones are food remains of
ancient hunters and cattle herdsmen who lived in pile dwellings along the
lake shores. From the vicinity of such structural remains at the Zanga
source two pairs of large antlers of red deer were collected and
transferred to the Caucasian Museum (Bayern, 1871). Pile structures were
also found near Novo-Bayazet and near Lake Gilli (Lalayan, 1929).
In 1947-1948, when the level of the lake started to fall, Dal' (1950b)
collected numerous bone fragments on the exposed part of the northern
shelf of the lake. The material included bones of deer, European wildcat,
boar and bear. A large collection of fragments of antlers and skullfragments
of boars and turs was obtained from a 1.5-m depth in the shallow-water
bay in the Artanishskii Gulf.
Material from the gray river sands of the Sevan coast (the lower reaches
of the Gavaraget River, near the Agbulakh rivulet and Cape Sary-Kai)
appeared to be older; it included fragments of antlers, fragments of skulls
and vertebrae of large bulls, similar in size to Bos primigenius and
bison. A fragment of skull of a dwarftur (Bos cf. minutus) was
collected from the same sediments near Cape Sary-Kai (Dal', 1950a).
218
At our request Khaveson collected some bones оп the Lake Sevan shores,
mostly on the northwestern shore near Shordzha, and on the southern shore
near Cape Noraduz and Sary-Kai.
222 We studied both collections. Most of the fragments in Dal's collection
are brown and light brown in color; in Khaveson's collection the material
is mostly of a light color. The bones seem heavy and permineralized,
which, however, is only due to the limey deposit on the surface. Bones
from the surf zone are strongly rounded and beige in color; those from the
sediments in quiet water and from silt are complete and of a dark color.
The deer remains mainly consist of antlers, often in a state of decomposition
due to their exposure to water and subsequent weathering on the shore.
Complete metapodia and jaws of domestic bulls and buffalos have been found.
From the degree of coloration and loss of organic matter the material can
be subdivided into two age-groups. The species identified and their
proportions are given in Table 55.
TABLE 55, Species and number of bones of mammals from Lake Sevan shores
Domestic
Па 69 ба а В ое вы В оо а 12 8 20
(О АО р а о у 1 — il
BOSS q AAC а ооо Ghaken Ate 61 23 84
ес, ое 6 2 8
Gael О gos ced kore Ag о о 3 3 6
Lae ERE о обоо бобов sous оо бов 70 67 137
ВИА ооо ин, Ц 5 2 7
Small cattle (no closer identification) ... 13 14 27
Subtotal 290
Wild
Vulpes vulpesh.:. . Е 2
тощо ate OSM 2.2.2 4). ее 1
Miedes ime les... оо. Ни 1
SiS SIGH OL AY ie ой. Пе ня 5
Conus ЭЛАР с о овоббоооросво 52
Subtotal 61
Total 351
The occurrence of dromedary is particularly interesting. The state of
preservation of the forearm, scapula and mandible (Figure 103), which
are colored light brown to a depth of 4-5 mm, indicates that the bones are
quite old (3,500-4,000 years).
Khaveson (1954b), on the basis of teeth indexes, identified the jaw
of the camel as belonging to а wild form —Camelus dromedarius
dahli. Confirmation of this identification based on more complete material
219
223
would indicate an extensive development of хеготогр с landscapes of the
Armenian Highland at some early stage of the Holocene.
It seems very doubtful that wild camels could migrate into a mesophytic
landscape of the present upland type in the vicinity of Lake Sevan. Other
wild forms (bear, boar and deer) are more characteristic of the mesophytic,
i.e., meadow and forest, environment than of the upland steppe and desert.
Material from burials and settlements. It has been possible
to date more accurately the bones from burial grounds and ancient towns
on the Lake Sevan shores, the Karabakh plateau, the Zanga shores, Debed-
Chai ravine and the upper reaches of the Khram, near Tsalka on the
Trialet ridge.
According to Piotrovskii (1949, p.92), the oldest Holocene site is the
Eneolithic, on the bank of the Zanga River near Shengavit, which
was excavated by Baiburtyan in 1936-1938. The site was dated as second half of
the third millennium B.C. Dal' has identified inthe site material bones of wild
goats, horses and fish; the horses were probably wild varieties.
Remarkable material, including antlers of red deer, was collected by
Kuftin (1939) in 1930 from Eneolithic (third millennium B.C.) mound burials
on the Trialet ridge in the upper reaches of the Khram.
A lower molar of abear (Ursus arctos meridionalis) was
collected by Kuftin from an Upper Eneolithic single burial near the village
of Ozny in the Tsalka area.
FIGURE 103. Lower jaw of Camelus cf, dromedarius from
the shores of Lake Sevan
More recent burials from the middle of the second millennium B.C.
contained fragments of skulls, metapodia and phalanges of giant bulls,
nearly the size of Bos primigenius, аз well as bones of dogs, horses,
boars, sheep and goats. АП the bones are stained with humus; they are
dark brown in color, and partially decayed. Kuftin thought that venerated
human remains were carried on bull-drawn carts to the grave site. The
bulls were then slaughtered and eaten, and the skins, skulls and limb bones
were buried with the corpse. These mounds yielded a silver bucket
depicting red deer and chamois and a gold goblet depicting lions.
1704 220
224
The same type of burial with remains of a bull was found in 1948 in
Kirovakan (Piotrovskii, 1950). The skull, better preserved than the
one at the Trialet locality, indicates that the primitive tribes who
built burial sites of this type had begun the domestication of Bos
primigenius. A pair of roaring lions, similar in style to Hittite
drawings, are depictedon a gold cup found in the grave.
It is known from Resler's (1896) descriptions ''of the sites of huge fires"
that "gre at accumulations of ashes and burnt animal bones — horses, rams,
dogs, boars, buffalos, and bulls — '' occur in the mounds and burial sites at
Archadzor near the village of Khodzhala on the Karabakh Plateau. The
presently accepted date of these monuments, which are related to the
Assyrian culture, is early first millennium B.C. (Piotrovskii, 1949).
Skulls of horses, bones of rams, camels and wild animals were found in
the dolmens excavated by Belek and Ivanovskii in Karabakh and near Kedabek
(Sysoev, 1925).
FIGURE 104,
Skulls: 1—Vulpes vulpes; 2—Martes foina; 3—Lepus europaeus
from Urartu burials on the shores of Lake Sevan
221
The excavations of the Urartu fortress Tsovinar on the southern shore
of Lake Sevan (Piotrovskii, 1944) revealed bones of domestic pigs, domestic
bulls and sheep. Large Urartu burial fields on the eastern shore of Lake
Sevan near the village of Zagalu were excavated in 1905-1908 by Lalayan
(1929). Big earth mounds of three types, which cover stone mausoleums
used for common burials, were found there and near the villages of Nor-
Bayazet and Adiaman. By the transitional Bronze-to-Iron-Age material
found there, these burials have been dated as early first millennium B.C.
225 (Piotrovskii, 1949, p.86). The sites contained skulls of martens (Martes
latifrons = Martes foina)and badgers (Meles urartuorum =
Meles meles minor) (Satunin, 1907b). From the same sites,
Shelkovnikov in 1930 collected a mandible of a horse, a skull of a young
pig, two skulls of foxes (Vulpes vulpes), four skulls of martens
(Martes foina) andaskull of ahare (Lepus europaeus), all of
which he donated to the ZIN.
The burial of the bodies of wild and domestic animals was of ritualistic
importance to the Urartus. Lalayan mentions that he invariably found
skeletons of horses, large cattle, sheep and goats in Urartu graves which
he disposed of at the site. It is not known whether complete bodies of wild
animals or only their skulls were buried (Figure 104).
TABLE 56, Species and number of mammal bones from Urartu fortress Teishebaini
Number
of
individuals
Number
of
individuals
Species Species
Domestic Marge si Ро ее. 1
Ре! пуле в... 1
о Omoaoo o timed dc I у й .
С Cefu lus арта о из. ... 2
hot oon ое DODO Um > bo ac
i Мера о ОВ р cosy << 2
ооо о оО: аи у
р. Apodemus Sylvaticeus,.... . 4
ООО cidkate: ОА РО у
8 ето тес ар а. ие а 1
Large Cattle: |... м Ви
ap wal ae gage PLUS: и, bas Raye “s 1
GEM) о о A Chor oO Обо Pct > an
ON US SSM GI. heer ЖЖ 1
SHEEP® оон ce. oo нь
Gazella subgutturosa..... 2
GOAL) aie silo соо о а ем
Соус... 4
© wees On 4 в Що 2
OM, ae. hs) ма 98 в
ооо Ч
ee a се тая
6 © ye Of Axe Ve
Ornaments from the graves usually depict red deer, A running bear
is engraved on a sard seal collected in one of the graves.
Excavations of the Urartu fortress Karmir-Blur (Teishebaini)
on the Zanga River near Yerevan (Piotrovskii, 1944, 1949) produced bone
material which reflects some features of the animal husbandry and the
faunal peculiarities of the Armenian Highland in the middle of the first
millennium B.C.
222
226
Teishebaini, an important administrative center of the Urartus, was
conquered inthe 7th century B.C. by the Scythians who invaded
Transcaucasia. Domestic and wild animals and various objects were buried
under falling roofs and walls as the fortress burned. Dal' (1952), who has
studied 4,667 bones from the Teishebaini excavations, listed the species
given in Table 56.
From his knowledge of the conditions of the burial and his personal
observations at the excavation site, Dal' concludes that gerbil and long-
eared hedgehog migrated to the ruins of the fortress at a later time. The
stone marten (Martes foina) andthe spotted cat (Felis lybica)
might have been domesticated species which were used to control rodents.
White -toothed shrew, hamster, field and house mice belong to the local
synanthropic forms which at that time invaded human dwellings. Bones and
objects made of horns of wild goat, ram, deer and goitered gazelle give
some idea of the hunting traditions in the northern part of Urartu. This
incidence of goitered gazelle bones is the first documented evidence of the
occurrence of this gazelle in the middle section of the Araks valley.
Material from shelters of predators. New paleontological
materials of the postglacial epoch were found by zoologists during studies
of accumulated food remains of predators in the caves and smaller caverns
of the Lesser Caucasus upland.
Polyakov (1882), in his survey of the caves along the shores of Lake
Sevan in 1879, found a skull of a weasel, jaws of voles, mice and ducks in
а cave at Mount Peschanaya Gor.
In his study of the present vertebrate fauna of the Saraibulakh (Urtskii)
ridge, Dal' (1940b, 1954a) collected and identified the remains of animals
accumulated by preying eagle owls and wolves in four karst caves. (Some of
the bones belonged to animals which entered the cave and died there. ) The
richest material was collected in a cave in the third gorge from the west,
in the western part of the ridge, above the village of Shagaplu. The entrance
into the cave is on the right wall of the gorge, 11 m above the ravine floor
(1,750 та above sealevel). The combined length of the sections of the cave
which were studied is 41.5 m. The animal bones were scattered and were
mainly found in depressions and chambers, occurring in large numbers
in the clayey surface layer. In this cave, 1,850 bones, belonging to at least
660 individual reptiles, birds and mammals, have been collected.
The state of preservation and the age of the bones vary. Some bones of
domestic animals which remained on the surface were better preserved
than those which were embedded in the ground. The bones are grayish
yellow in color. Calcareous crusts up to 1.3 mm thick and stalagmites up
to 5 mm high formed on some of the bones. The oldest bones dissolve
completely in hydrochloric acid. Па! estimates their age from 2,000-
4,000 years. They include bones of jackal, domestic dog, fox, stone
marten, tiger polecat, badger, hare, ass, horse, deer, bull, wild goat
and Armenian mouflon.
We have studied the skulls of badgers from this cave (Vereshchagin,
1951b) and found them to be closer to that of the Urartu form of badger than
those from the Sevan burials. This fact makes it seem possible that some
of the bones are older than the age assigned to them by Dal'. The jaws
of a large pika, found in the far corner of the cave, are among the oldest
bones. The remains of white-toothed shrews, Myotis, foxes, weasels,
223
227
martens and rodents are of modern types. Most of them were washed out
of the pellets of eagle owls by water.
The identification of the material from the four caves resulted in the long
list of species given in Table-57. Bones of larks, chough, pigeons,
Hungarian and rock partridges, Ophisaurus apus and lake frogs were
also found.
The location of the caves in a transitional landscape zone and the ways
by which the animal remains were introduced into them preclude their
usefulness as a basis for any far-reaching conclusions on changes in the
climate and landscapes during the Holocene.
The list of wild species of the Saraibulakh ridge shows that large species
(jackal, bear, pika, kulan and deer) do not occur there in the present;
their disappearance can undoubtedly be attributed to man.
TABLE 57, Species and number of mammal bones from caves on the Saraibulakh (Urtskii) ridge
Number
of bones
Number
Speci
Ae cae of bones
Species
Domestic
ID IS (Sty roe БОИ, CE ARC ON, Rah mee Mien cictarmtedlionss ее. esc 14
HOBO We). Ю.И. oo BY A, Fe ThistDyibiioalt 20. FUER ке A i
J ACIOR Dre teat ее рее ое И Lepussemropiaien(sa.s.% ления, 98
batgeveattlen к аль. aril - Oicihiait Gniaygspagts sess bee МЫ 3
А mc cer coe ров Allactaga м тать. 9
ОА, ое А Mie iO eS рее ева 135
Cricetulus migratorius 135
re oe a MES Ger TG emus allie t Ws a, conics 794
ан А SOCMAMUNS costes cts a ee 140
Mi My atl ins: Rar. ee at М Ма 6
BVINAGCeUSIeUTOpB ens я Lh. 44. Aryilcw laAMLeDresitrists. mt а 2
Cro cidu ray Tussi) asp, wz). Ме Blo bars” we es/clense tiie aesepai 323
Myotis oxygnathus......... Equus hemionus,....... erg 4
Са aT CUS oon) epaysneue Vousn one GClervyuis арии о 24
нок Ее Е ПВО Ga Pa Ae А а не Е: 85
МЕ У ее Ovis gmelini 76
оо erento rao rate
Ме ЧИНА a Se Ne ae ete ee Pe Ome Pe AMO IDIOMA PT te. es
Мате Seto iia te tate . Rh ne tel et etre
Vormela peregusna ,
Taken аз a whole, the faunal complex of the region, even in the oldest
times, was very close to the present faunal complex, which is characteristic
of the arid plateau landscape of southwest Asia.
In southeastern and central Armenia, Sosnikhina (1947, 1948) did a
thorough study in 1946-1949 of bones from eagle owls' pellets in three
different altitudinal vegetation zones and landscapes. These were: 1) zone
of spiny tragacanth formations (1,500-1,600 m above sea level); 2) zone
of upland fescue steppe and arid thin forests (2,100-2,200 m above sea level)
in the Azizbekov area and near Lake Sevan; 3) zone оё mountain meadows
224
(2,600 m above sea level) in Nakhichevan A.S.S.R. The material, consisting
of several thousand bones, illustrates very well the distribution of small
mammals on the high plateaus and the effects which changes in climate
and landscape can have on formation of ecological assemblages. These
data are summarized in Table 58.
Most of the bones studied probably accumulated during the last centuries,
possibly only during the last decades.
The two main assemblages of small mammals, the xerophilous and the
mesophilous, either replace or supplement each other, reflecting in part
the landscape of the region and in part the distribution and hunting behavior
of eagle owls.
These data also indicate some etiological features of the behavior of
eagle owls: the length of time they occupy their shelters, their habits of
preying on a wide variety of species, choosing their prey in various
biotopes, and leaving their pellets near the hunting ground. Inthe winter some
of the eagle owls probably migrate from the highlands to the valleys, and ,
228 in the absence of shelters on the hunting grounds, they may migrate to
another area to find day resting places.
TABLE 58. Species and relative abundance (in %) of mammalian bones in pellets of eagle owls of
Armenian Plateau
Chirakhan Southern slope : у Slopes around
i : Dzhagri ravine ;
Species ravine (1,500- | of Sevan ridge Conn аъоме Lake Aknalich
1,600 m above | (2,000-2,100 т |`’ (2, 60011 above
sea level) above sea level) ео sea level)
Erinaceus europaeus ........ 2.4
Crocidura russula ........ =
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HOS ПИШИ 5 S55 5000 co0D00 =
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MSVOUIS OrxsyemawntS 5555000006 a
Mustela nivalis caucasica,., 1.2
Гери CUPROPACUS 5455006000006 7
Фет ар Ио ween Е 4,7
ИЕ eel Ween 55 oo G0 G6 р,
"Ооо И ОИ Е 6556566000 =
Apodemus sylvaticus........ ar
МАТИ G5 coanooos0000008 a
Cricetulus migratorius...... i
MESOCIULCSTWS ИМЕЯ 5 65's 65% 17.3
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ПИ, S@ GUIS Бовобоооав бов ов Ee
MPa Ore. оо 2.4
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Arvicola terrestris..... . aa
ВПП ИО so ag500006 2.4
225
229
230
The occurrence of pika bones is of interest, sinces this species has
probably become extinct in modern times on the Armenian Plateau.
In the upland semidesert of the Araks valley, bones from the pellets of
owls and eagle owls were collected by Sosnikhina (1950b) in a cave of the
volcanic Mount Karakhan (900 m above sea level), near the village of
Novyi Shakhvarut, west of Yerevan. In 1947 we collected material in caves
and under overhangs at the foot of the rock formations known as Alindzha-Dag,
Ilyanlu-Dag and Darry-Dag, and in caves near the village of Klit, east of
Ordubad, at the beginning of the Araks gorge.
The present-day landscape of the Abrakunis area consists of plateau-like
spurs, covered with spiny astragalus bushes, mountain saltwort and weakly
developed ephemeretum. The relict shrub vegetation, which had
persisted in the valleys, consists of almonds, buckthorn and ephedra. This
vegetation indicates that a xerophytic landscape existed in the region even
before cattle husbandry. The plateau-like spurs are cut by dry canyons with
rock-covered areas and caves, which are the resting sites of owls and
eagle owls. There has been a survey of one such cave in the ''Hyena ravine"
Kyaftar-Dara (Figure 105).
On the left wall of the Araks gorge we surveyed three karst caves located
between the Ordubad and Megri stations near the village of Klit.
These caves were visited in 1879 by Polyakov (1882) on his reconnaissance
of Paleolithic sites. In a trench in the anterior chamber of the middle cave,
at a depth of 1.5 m, Polyakov found two pelvic bones of wild goat (Capra
aegagrus), leg bones of European brown bear (Ursus arctos), several
bones of hare (Lepus europaeus), "femur of a small carnivore similar
to a dog'' and bones of a pigeon. The excavations revealed the following
sequence from topto bottom: a layer of flaky clay and excreta of bats, 35 cm
thick, clayey beds with excreta and bones of bats.
All three caves are located in a wall of dolomitized limestone on the
eastern slope of one of the small valleys in the Araks ravine. There are
other caves in this limestone which, however, are not easily accessible.
The biggest cave is the middle stalactitic cave ''Dlinnokryly'", described by
Satunin (1915a). The entrance slopes downward into a cavern, from the
middle of which the floor slopes upward to a long, narrow corridor with
calcitic encrustations. From the posterior part of the cavern another
twenty-meter-long corridor leads to the left into a lower chamber, 15 m
wide, and to yet another corridor, this one proceeding a length of 50 т to
a dead end. The combined length of the cavern and the long corridor is 250m.
The height of the first chamber is 12-18 m; some sections are higher.
In the talus left by Polyakov's excavations bones of Miniopterus
schreibersii were observed. Inthe left corner of the cave, near the
resting place of some carnivore (panther?), we found 14 gnawed bones
(horse, ass, sheep, tur) showing a recent mode of preservation, and
several bones of Miniopterus schreibersii covered with calcitic
crust. Bones of a young panther (Panthera pardus) and stone marten
(Martes foina), also showing a recent type of preservation, were found
in the stone-covered middle section of the cave.
Of the present cave inhabitants, we caught only one greater horseshoe
bat (Rhinolophus ferrum equinum) and counted only ten flying
Miniopterus. Satunin in 1893 counted Miniopterus inthe thousands.
Footprints cf panthers, martens and white-toothed shrews occur in the
ancient dust.
226
(229)
231
FIGURE 105. Entrance to cave inhabited by eagle owl in Kyaftar-Dara ravine
Photograph by author, 1947
Some rodent bones from the pellets of eagle owls were collected near
the mouths of the two marginal caves.
Comparison of Sosnikhina's collections of small species with ours
indicates a considerable increase — from west to east — in the xerophilous
fauna and a concomitant species impoverishment in the small-mammal
complexes (Table 59).
All of this material from the Arals valley — up to 1,500 bones from owl
pellets — was collected from the surface and generally shows a recent type
of preservation. It gives no indication of the appearance or the disappearance
of any species in the semideserts in our time. A similar composition of the
fauna was established through sampling with traps. However, in the
interests of accuracy, it is necessary to mention that a mouse-like hamster
was discovered as recently as 1938 on Mount Darry-Dag near Ilyanlu-Dag,
and that bones have been found of Asia Minor hamster which does not inhabit
the upland semidesert zone at present.
Тату: за EW ibwws jopland
No bones of the postglacial epoch are known from the northern foothills
of the Talysh Mountains and the Lenkoran Plain; neither are food remains
227
known from the Talysh highland steppe. However, ancient burial mounds,
in southern Mugan, as yet untouched by archaeologists, are evidence of
ancient human habitation of the region.
(230)
TABLE 59, Incidence of species (in %) in mammal remains from pellets of owls and eagle owls in caves
of the middle Araks Valley
Abrakunis,
Kyaftar-Dara
cave
Ordubad,
Dlinnokryly
cave
Novyi Shakhvarut,
Mt, Karakhan
Species
Erinaveus seu Top Meas ier «een
Hemiechinus: a Wig uses pees eee
Grocidura russ Wildy aa ee dio ae eas
Miniopterus schmembens tine. jen
Mustela miva 15 aver. ole an ais
Lepws €wrop a € US ее
Allaetaga ма evacuees nee
Allateta вает ее email
Mis) цессе
Apodiemius. sy lL vane но еее ee
Метгошез. Бас кет Е cae
М. Pie ES UCUIS: И ue © ее
Мезоесттсети$ .aAMEMISy ое
Cricetutus тол оваветииь S| 4. ее
Calomyscus bailwardi ...... ...
Атугсо1 а террасе о ors
MirGriotulsmarivallits’ . Steere ee ее
М soleil a Lis, зо ое gods
Edo bans, ре се ee
In 1945 I collected rodent bones in small caves inhabited by owls located
in three sections of upland steppe, covered with grass and spiny astragali,
in the upper reaches of the Vilyazh-Chai, near the Dymanskii post and in
the upper reaches of Vassaru-Chai, near Kelakhan and Kel'vyaz.
TABLE 60, Species and number of mammal bones from owl pellets in upland steppe of Talysh
Species
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CLUCETUNUSM та OTIS ena Rees № a vela te
Miesoernceitiisy aiming Goya. аа: ие
мМистоли зева Ба о-в
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Ea Mob а лв, im Ge SiGe т еее > cus а cee .
Е о ооо резво .
228
The age of these bones is not more than 200-300 years. In most cases,
they were not protected by soil and consequently were subject to weathering.
The combined data for all the three sections are given in Table 60.
All the small mammals listed still inhabit the region, forming the Recent
assemblage characteristic of the upland steppes of the northwestern Iranian
Plateau and the Armenian Highland. The list does not include the European
hedgehog, long-tailed shrew, steppe mouse and snow vole. These species
are very rare in the region. (Relict snow voles occur higher in the
mountains. ) The similarity of our list to Sosnikhina's (1947, 1948) list,
given for the tragacanth-covered steppe areas of the Armenian Highland,
proves a considerable degree of affinity between the faunas and between the
landscapes of the two regions.
On the adjacent Iranian Plateau, Coon (1951) collected Holocene
mammals from the eastern part of the Elburz Range, 50 km from Asterabad,
in Mesolithic and Neolithic beds of the Belt cave. Among the 1,170 fragments
of bones, 18 species have been identified. The most abundant is goitered
gazelle (Gazella subgutturosa, 67%), followed by Bos
Пена еа ис (18-89%) coat (арта 5р. 5.19), ram, (Onassesipe.
Фа рлаи seal (Eshioje aye as pile aw ice lo) andidopa(@anas sp, ,
1.7%). Also mentioned are individual occurrences of Vulpes sp., Felis
Sjoe, Wilweigieucleis, ‘Crp Oli Sos, (eS Oi нноею, Isl yas te ss) Soe 5
Mus sp., Equus asinus, Sus sp., Cervus elaphus and Cervidae.
232 Identification of the bones of the Viverridae, European beaver and Bos
primigenius needs to be confirmed. The remaining species are to some
extent similar to the present assemblages of mammals on the hot plateaus
of northwestern Iran and humid, wooded Asterabad.
These studies of the various types of Holocene deposits of the
Armenian Highland, Araks valley, and Talysh and El'brus highlands show
that a number of large animals (bison, Bos primigenius, kulan,
goitered gazelle) inhabited the region in the not-distant past. They also
reveal shifts in the areas of distribution of some smaller species (suslik
and Asia Minor hamster).
The results of the studies of Holocene localities on the Caucasian
Isthmus can be summarized as in Tables 62-63.
In our studies of the bone collections from the post-Paleolithic cultural
beds and from diluvial sediments, we were able to trace the appearance
of some species of large mammals in the Caucasus in postglacial time.
The species migrated to the Caucasus from adjacent areas. Extinction of
many of the Pleistocene species was also revealed by the studies. The
rapid development of animal husbandry and its characteristic regional
features, as well as the steady increase in the proportion of domestic over
wild species in food remains from the Neolithic to the present, could be
traced through the bone material. Studies of bone material from the day
resting places of eagle owls produced a more precise delineation between
past and present distribution of many insectivores and rodents. The rarity
of multilayered Holocene deposits of smallbones and the abundance of
younger bones in small caves and under rocky ledges might indicate a
discontinuity in the availability of shelters and in the accumulation of food
remains of owls. It is possible, however, that rapid mechanical weathering
229
233
of *he exposed rodent bones and the frequent collapse of small caves might
have created the appearance of such a discontinuity.
The studied Holocene material consists of 77 species of mammals which
inhabited the Caucasus at the end of the Upper Paleolithic, some of which
became extinct toward the 20th century.
Comparison of the lists of the Holocene and Pleistocene species indicates
pronounced changes in the compositions and distribution of the mammalian
fauna of the Caucasus, occurring mainly at the Pleistocene-Holocene
boundary (Table 61).
Thus, the Holocene in the Caucasus is characterized by the complete
disappearance of proboscideans* and apes while perissodactyls and
artiodactyls decreased catastrophically. The increase in the number of
rodents in the Holocene beds is accounted for by the differences between
excavated and open sites.
TABLE 61, Number of mammalian species from the Pleistocene and the Holocene in the Caucasus
Pleistocene
Species Holocene
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Reliable conclusions оп the appearance or disappearance of species in
the Holocene can be drawn only with respect to the mass species. Overall,
the species complexes and locations of the Holocene deposits exhibit a nearly
modern faunal composition and distribution pattern.
A review of paleontological material on post-Pleistocene fauna shows
the following changes:
Insectivora — At least five species crossed the Pleistocene -postglacial
boundary: mole, white-toothed shrew, water shrew, long-eared and
European hedgehog. In addition, three species of white-toothed shrew
(Crocidura suaveolens Рай, Crocidura leucodon Hermes,
Suncus etruscus) occur inthe Holocene; these species probably inhabited
the Caucasus in the Pleistocene also.
Chiroptera — Greater horseshoe bat (Rhinolophus) is known from
the Pleistocene beds. The Holocene beds in the caves of the Lesser
Caucasus uplands commonly contain bones of Myotis and Miniopterus.
Primates — There are no data on the occurrence of Holocene macacas.
* In view of the finds of geologically young mammoths in Pyatigor'e andin the Gori depression, the question
of their possible survival through the Pleistocene in the Caucasus deserves further investigation and requires
additional corroborative material,
230
244
Carnivora — Of the 21 species known in the Pleistocene, only 15 have
been recorded in the Holocene. In the Holocene of the Caucasus there are
no steppe or cave bear, cavehyena, Panthera leo or cheetah and probably
no glutton. Jackal and striped hyena occur for the first time in the Holocene.
It seems possible that at this period a number of other southern carnivores
(jungle cat, lion and tiger) migrated to the Caucasus although their remains
have not yet been found. The absence of lion, tiger and cheetah bones from
postglacial deposits canbe accounted for by their scarcity in the historical
epoch.
Lagomorpha — European hare is the only species known from the
Pleistocene. The Holocene deposits contain remains of European hare and
large pika which probably existed in the Lesser Caucasus in the Pleistocene.
Rodentia — A number of important changes in the composition and
distribution of the rodent fauna occurred during the Holocene. The porcupine
andthe Apsheron vole became extinct; the Asia Minor hamster retreated
from the Apsheron and from Imeretia; the areas of distribution of the suslik
of Asia Minor on the Armenian Highland and of the Caucasian snow vole in
Ciscaucasia decreased. The finds of remains of black rats in Neolithic beds
on the Black Sea coast and in Kura Plain beds of the middle of the first
millennium B.C. are a basis for assuming that the species was autochthonous
and that it survived through the Pleistocene on the warm, humid lowlands
near the Black and southern Caspian seas. A similar conclusion, if applied
to the Norway rat, would be oven to question. The occurrence of European
beaver in Neolithic times in Colchis has been confirmed.
Most of the other findings fill in details of the distribution of extant
species.
Perissodactyla — The materials studied give no indication that the woolly
rhinoceros and the Equus hemionus survived in the Caucasus through
the Pleistocene. The occurrence of the modern kulan in the Holocene of
eastern Trans- and Ciscaucasia has been recorded. There were undoubtedly
wild horses in the Holocene, but it has not yet been possible to distinguish
their bones from the bones of domestic horses of various origins. Wild and
domestic horses were the main animals used in sacrifices and for food by
the Caucasian tribes during most of the historical epoch.
Artiodactyla — Boar were widely distributed in the Caucasus. In the
Holocene the species was one of the primary game animals. Bos
primigenius were also abundant in the Lesser Caucasus. So far there
is no evidence of the existence of wild camels in the Caucasus in the
Holocene.
The wide distribution of red deer has been traced over nearly the entire
Caucasian Isthmus from the Pleistocene to the 19th century. Roe deer
and elk inhabited the lower Don valley, the river valley forests and reed
jungle of the Kuban and the wooded lowlands of Ciscaucasia and Colchis to
the end of the 18th century. No Holocene remains of Cervus
pliotarandoides and fallow deer are known.
The occurrence of bison has been established for the Bronze Age in
the Lesser Caucasus, and for the Middle Ages of the present era on the
middle and lower Don. Saiga disappeared from eastern Transcaucasia in
the Lower Holocene. Remains of goitered gazelle begin to occur in beds
dated second millennium B.C. in eastern Cis- and Transcaucasia. They
are particularly abundant in the Apsheron Peninsula beds of the Middle Ages.
231
234 TABLE 62. Species and stratigraphic location of mammal bones from the Quaternary of the Caucasus
235
Orders and species
Insectivora
Erinaceus europaeus Г. ($. lato)
Hemiechinus auritus Gmel. (s, lato)
SOP TURAL HL tery ests tel sees ss
ЗОО TE bs oh Pe EAR OE, Е
Neomys fodiens Schr, ($. 1ато)....
Crocidura Lewcodon Herm Je. i
Coa russuda Pali (s. ato) i sss Bae
SUMS WS вас (Sa Vie рее ode
IPM cy OEUNOEI NOE Sele 5 3 Hos ooo
6 SD) CRIs Or ceaidisiedioe Ora
Chiroptera
Rhinolophus ferrum-equinum Schr.
Myotis oxygnathus Mont, ......
Miniopterus schreibersii КИШ...
EMITOpterakGelaron . он
Primates
DCMCRAICAMS Dp choke eure ac aii eu ey, = Supers
Carmivora
CAMS sae И atte reyes) a ONO Cae
а: Sagal EO) haa ae ie a ene
С. lupus apscheronicus М. Ver,
“GC. tamamensis М, Мег. 111% 38
Сам 8ря phe cc па ВЕ Е
Слои
Уре со та ера AES -mebey. eye ie на
У. унирес т. врал
У. vulpes aff, alpherakyi Sat. ...
WS WSS ANC воз ee pcre Че hel
О. arctos binagadensis М. Ver...
WU. speliaeus Rosm (sy lato)c G2 „Г...
U. ressiten иво: ee.
Hy aena shy vena. оу
Crocitta «ретаеа соч io awe
Mines, om nids вх. ms ии
М. боди а: slairifironisesai.y eet В
Мое ГОША: ео а ел
Ne Е ВЯ, Арба ЗУВЕ on ceed,
Vormela регеризпа Guld. ......
Lujtramihut ral 19:2. Sts де
Mustela nivalis L. (5. lato) .....
* Species of the transitional Upper Pliocene-Lower Pleistocene period on
with an asterisk.
Lower
Pleistocene
Middle
232
Upper
Mesolithic
Holocene
o Ей Е
$ 50 50 {>
= < v. < `—
Ф Ф Е
= N < = 5
[>) я S ne} 2
oO о о к [9
[= -
7. a) 4 = 5.
+ + +
— + +
— — +
— = +
== + +
— + +
- + +
_ + +
+ + +
— = =
— - +
+ + +
= se =
= + +
os == =
+ = =
a + +
т af A:
os = ag
+ = be,
the Taman Peninsula are marked
TABLE 62 (continued)
Orders and species
MELISS WEISS № (Gs Ею) Ses 56 o 4
Mp, DNEILES MARLOW ЗЕ 5 sno 5
М. MmETSS в, AMO SA 52564000
MCIES S06 чобы Bid o-oo 4 ао оо
Gulo aff. gulo L,
Felis silvestris Schr, ($. lato)
I's WWIDLGA 95 goog os50509006
Po аш. WVICAa 163% oooacaodocg
Во ухе о (Sip LAO) о ctr ake orem
Panthera pardus L,
Р. spelaea Goldf,
Рама G0 oooaa
Ce ee
оон
ооо
ACHMOMIPS Elis MIDAS бе 55 55
Lagomorpha
OCMOLOMA Во бсосовобовосасов
LEPUS GUROPACWS Pall, о ръоовос
L, europaeus gureevi I, Grom...
L. cf. europaeus
Rodentia
МА О EE SP ob aodguogues 02000
те 113 рузмаець Раш. ooo. «
С. aff. citellus xanthoprimnus
WOM, coors oo0gsoga000020000
CASEOE WiDEH by собососбосочдо
* С. tamanensis №. Ver.
* Trogontherium cuvieri Fisch,
Hystrix cf. leucura Sykes ......
H. vinogradovi Arg,
Dyromys nitedula Pall,
ИИ yy Wits ho бро 6 G0 Glo dina од
subtilis Pall,
Оо ©, ФЕИ VAM, Saves ade ai
Allactaga jaculus Pall,
А. jaculus bogatschevi Arg.
ЕО, eNOS, mike ie sis о ро ао Е
Sicista
ве ие Е
ев =
. СЁ. williamsi
iN EME WET ПО Yn G6 ood 000 0100 0
Spalax microphthalmus Guld.
So В ING 5 Soon a0 oo
Micromys minutus Рай.
Mus musculus Г. ($. lato)
Apodemus agrarius Pall.
A.sylvaticus Г. (s. lato)
ROEM) MANIC Ibs 9 6 6 oe no oo
R. norvegicus Berk.
see ewe
. Williamsi dzhafarovi I. Grom.
+b +
Pleistocene
= 5
м)
jou
Ss a
= =)
+ +
a Е
= +
+ +
£ a
= +
+ +
+ +
a =
= +
-|- —
Зы —
+ +
— +
+ +
и oa
+ =
— +
г ==
ws 5
ek
a. oe
+ a
г he
233
о
=
=
=
о
n
o
=
=
Neolithic
+
Holocene
9)
50
и
<
|=)
5 5
с [=
т
д a
= nie
fe me
— +
+ +
— +
bu fie:
— +
— +
ar +
+ +
+ +
+ +
+
|
|
Middle Ages
+
Present time
236 TABLE 62 (continued )
Orders and species
Mesocricetus
Mi ат: araltiise Weiter. Bt orc. Е
Me aura tus Tadic em Nehine sistant Е
Maura tuls ра ео а Argyl nan kot. ae
CTUCE оао а vO а» Pauly ie ec
С. migratorius argiropuloi I. Grom.
Calomyscus bailwardi Thos,
Meriones erythrourus Gray, .......
М. erythrourus intermedius I, Grom,
MiP DLAe Rr ОНО: де аа steno © he hehe
Me persaicuis „Вами. о са aha ote
Ва ити о ас мхи AEEUILS NG ое
Arvicola tentestinis №. ©. Пато) jb. |
Mie rons: anvialus: Paley, be tia sates ie
М. socialis Pall,
. ef, roberti Thos,
nivalis Mart,
aff. gud Sat, ,
A ЕО MOS, (8 ts, dick mie wht amie od
apscheronicus Arg,
Prometheomys aff, schaposchnikovi
За,
Ellobius talpinus Pall,
Е. LU besice ns TOs. йа.
Е. aff. lutescens Thos,
Muridae реп.
Эа ‘ares sx
<
Мия, UTES ЗИ АЕ eas fot аа: ное ое
м.
м.
м.
щенки
Бот Saree) |B
TC CATE CR ЛИС ORDER CON, RCH? (hes a
Proboscidea
*Elephas meridivonalis Nesti 2. % 2.
AE Тала, AUC ил дор в
В trogomet he riyeP onl) и:
Ирана цели BUNT 5 si
Perissodactyla
Equus. цешопи Ра, ет
Е. aff. hidruntinus Вер. .,
Е, caballws Г. ЦТ, р ae oie be
Е. caballus gmelini Ant,
Ve! vue, чае
*E, aff. sissenbornensis Wist.
SE, Si KeTLOM US, Cocchi) оо dene aie
Rhinoceros tichorhinus Fisch.
Вели mete kis [aegery opis oH ane Uae
Rh. binagadensis Dzhaf.
Rh. sp.
Lower
+ +
Pleistocene
Middle
234
Upper
о
Me!
=
=
Е)
п
v
>
Neolithic
Holocene
vo
50
ча! ©
к <
=
5 5
= oq
fon) —
+
— +
+ +
_ +
— -
+ +
+ +
+ +
— +
+ +
+ +
Middle Ages
+
+++
+
+
Present time
+
+ +
+
distinguished from domestic
horses
237 TABLE 62 (сопипиеа)
Orders and species
*Rh.cf. etruscus
Elasmotherium
Se CAMeCASTE WGN Зо 4 op on 0 4
IPE Covey NET te ot AeA ten at
sibiricum Fisch. .
Artiodactyla
Эйс SevTOKea №. TAI) aacenoace
5. apscheronicus Bur, et Dzhaf,
аи а мещане IN, WEE 556 aoe oa 6
ali poyyworewiwsy Ho so og oan aes
Camelus dromedarius dahli eee
CiGIIISD)” cio alaradian oo ce gid
С. knoblochi Nehr, ор оросо <
*Paracamelus gigas Schloss. ..
Cervus elaphus L, (s. lato) ...
*Eucladocerus sp. ,
VCGIMIGAS BMG “ods oage 50a t
eee ee
*р. cf. kujalnikensis Chom.....
Dama cf, реет а Brooke
Megaceros euryceros Aldr, ,.
*“Megaceros sp....
Alces alces caucasicus N. Ver.
Mo Big В© 08 Iho оооввоовоевь
ооо ee ee
ааа се Ss isl cog ood aoe
Capreolus capreolus КГ. ($, lato)
SAGA CAC etes by 65000
see ee
Gazella subgutturosa Guld, ...
PAGE AGIMNEV SON Malai! og оные
*Tragelaphus sp.
*Strepsicerotini gen, et sp.
Rupicapra rupicapra caucasica
Аа
Capra cylindricornis Blyth ..
©. CGAMECASTOR Guill, рой ва
Co Aw, с аще аще а И
Co веваетиз, (СЕ 65 6 bleale oe o
OwviLs acini wibnins 65686 5 o\ba0 oc
Gl, Gi, РАЗОМ МКИ 556504540
С: ma mimosa cise. cls Per eee
BiOsS) ИИ ИЕ wrestle a) ens) is ao
В ем 5 о ово nl ole
В. ENO MOE TOS иле и.
В тата да еее ее
Bison bonasus caucasicus Sat,
В. priscus Boj. ($. lato)
“Ben Che SiG MO tems Kis теч sree cites
Ес eet Sennen sternite
“BL ОТО AGN E OSs teen ee м
o
- —
o x)
= 55)
2 =
3 =
es oe
= +
ci we
+ +
+
= +
ыы =
A =
= +
+ =
4 aes
= +
+ +
Е =
a: =
+ +
= +
+ ee
i =
a =
+ +
+ +
— +
= +
+ +
a ыы
+ +
235
Pleistocene
о
‘=
=
a
[=]
nan
‘5
=
Neolithic
+ + |
Holocene
oO
oO
тЫ
а <
=
5 5
FA a
aa) any
+ +
+ +
sa at
+ +
+ +
+ +
+ +
— +
A es
+ =
Middle Ages
Present time
238 TABLE 63. Stratigraphic summary of main localities of Quaternary mammalian index species in the
Black Sea region
Subdivision of | Marine
Quaternary beds Western Ciscatcasia
and Russian Plain
Climate and landscape
from the composition
of mammalian fauna
Western Transcaucasia
Sarkel fortress, Recent climate and
9-13th centuries
European brown bear
European beaver
kulan
elk
red deer
saiga
bison
CI- 284
increasing influence
of man on landscape
Present time
Semibratnoe site at
Kuban estuary
wolf
otter
boar
deer
Development of broad-
leaf forest in foothills
Iron Age
Huts of Colchidians at
Ф
a
в: = Anaklia and Ochamchiri
‘, 5 jackal
5 Е = boar
о o
я 2 roe deer
[2] Е
= o red deer
о
uv
[24
Mud-hut settlement at
Gelendzhik
lynx
hare
red deer
dolphin
CI-417
Neolithic
Akhshtytskaya cave,
upper beds
European brown bear
badger
lack rat
roe deer
CI- 460
Strong warming and
development of steppes
in foothills; waning of
valley glaciers
Mesolithic
* Calcination indexes of bones (СТ) by collagen analysis (Pidoplicnko, 1952); analyses aone in I.G.
236
239 Caucasus and adjoining lands
Caspian region
Marine
beds Eastern Ciscaucasia and
Russian Plain
Dzuars of North Ossetia
15-19th centuries
red deer
roe deer
Caucasian elk
east Caucasian goat
chamois
Caucasian bison
Settlements of Sunzha
valley, Isti-Su, Alkhan-
Kala
kulan
red deer
5 Dzhemikent
5 fox
2 bear
я kulan
2 goitered gazelle
Cl- 275
Sosruko grotto on the
Baksan
hare
little suslik
boar
red deer
Caucasian goat
CI- 241
Chokh site in Dagestan
hamster
mouflon
CI-317
Pidoplichko's laboratory in Kiev.
Settlements near Baku,
Eastern Transcaucasia
9-13th centuries
fox
kulan
goitered gazelle
seal
and settlement at Sumgait
striped hyena
black rat
boar
ted deer
goitered gazelle
237
Lesser Caucasus upland and
northern Iranian Plateau
Settlements at Kayakent and | Burials in jugs at Mingechaur |Karmir-Blur fortress
stone marten
Armenian mouflon
wild goat
goitered gazelle
Pile dwellings and burials on
Sevan shores
stone marten
badger
boar
red deer
dwarf tur
CI- 207
Belt cave at Asterabad
porcupine
red deer
goitered gazelle
Bos primigenius
seal
240 TABLE 63 (continued)
Black Sea region
Subdivision of | Marine
Quaternary beds
о
aS
Go
=
az
vo
= [>]
| Я
[5% з =}
5 ex |
te О
Ф [>]
5. o
7.
=
№
os
bo
я
a)
i
o 9)
г м
о
|2]
~
”n
.[ч
vo
—
о.
Ф
о
=
п
+
3
—
=
J
N
=
о
_
S|
=
=
wo) Ф
|9 Е
С.) х
BS 5. 5
= o =
et —
и =) я
ae) vo
As о
2 =
<
Western Ciscaucasia
and Russian Plain
Bone-bearing alluvium
of late terraces of
Don
fox
small pika
little suslik
great jerboa
water vole
horse
elk
red deer
reindeer
saiga
Il'skaya site at Krasnodar
cave hyena
mammoth
boar
red deer
saiga
Bison priscus
CI-518
Bone-bearing alluvium
of early terraces of
Don
Siberian polecat
little suslik
jerboa
Allactaga sibir
ca saltator
Eversmann
Spalax
red deer
i-
238
Climate and landscape
from the composition
of mammalian fauna
Western Transcaucasia
Gvardzhilas cave General strong cooling
mole and probable displace-
European brown bear ment downslope of
glutton landscape zones
Asia Minor hamster
Prometheomys
Caucasian goat
bison
CI-512
following glaciation of
high ridges
Sakazhia, Devis-Khvreli,
Mgvimevi caves
cave bear
horse Major uplift of the
European beaver mountain ranges
elk
chamois
Caucasian goat
argali
CI-580
Altitudinal zonation of
modern type
Akhshtyrskaya cave,
lower beds
wolf
cave bear
common hamster
red deer
giant deer
Bison priscus
CI- 706
241
Marine
beds
Khvalynsk stage
Khazar stage
Eastern Ciscaucasia and
Russian Plain
Diluvium in Yankul' gulley
on upper Kalaus
horse
saiga
Upper travertine complex of
Мг. Mashuk in Pyatigor'e
mammoth
horse
roe deer
Bone-bearing sands of Terek,
terraces at Mozdok
Bison priscus
Bone-bearing alluvium of
ancient extended Volga
valley
cave hyena
Panthera leo
mammoth
woolly rhinoceros
horse
Camelus knoblochi
giant deer
long-horned bison
СТ-484
Caspian region
Eastern Transcaucasia
Bituminous loams Kir-Maku
and Artem I,
wolf
horse
red deer
saiga
European tur
СТ 158-208
Bituminous sands at Binagady
on Apsheron Peninsula
long-eared hedgehog
corsac fox
Ursus arctos binaga-
densis
tiger polecat
cheetah
red-tailed gerbil
porcupine
Rhinoceros binaga-
densis
red deer
saiga
tur
CI 450-520
239
Lesser Caucasus upland and
northern Iranian Plateau
Diluvial loams at Maragheh
wolf
cave hyena
horse
rhinoceros
argali
Tamtama and Bisotun caves in
northern Iran
jackal
cave hyena
panther
gerbil
ted deer
goitered gazelle
Zurtaketi site on upper Khram
horse
ass
mouflon
Bison priscus
242 TABLE 63 (continued)
Subdivision of
Quaternary
Pleistocene
Pliocene
Lower Paleolithic
Eolithic
Chauda beds
Krasnodar beds
Black Sea region
Western Ciscaucasia
and Russian Plain
Bone-bearing sands at
Girei and Krasnodar,
middle beds
Elephas trogon-
therii
long-horned bison
bear
Bone~bearing sands at
Girei andIvanovskaya,
on Kuban, lower beds
southern elephant
Elephas antiquus
deer
bison
Conglomerates and sands
at Kuchugury, Sinyaya
gulley and Tsimbal
on Taman Peninsula
Trogontherium
cuvieri
Castor tamanen-
sis
southern elephant
Elephas antiquus
Elasmotherium
caucasicum
horse
Cervus pliota-
randoides
camel
antelop
bison
CI 913-1203
Western Transcaucasia
Kudaro cave on upper
Rion, lower beds
macaca
Siberian red dog wolf
cave bear
panther
Asia Minor hamster
marmot
Prometheomys
vole
porcupine
rhinoceros
boar
roe deer
argali
С1- 914
Fauna unknown
Sands and gravels of ancient
terraces of Kuban at
Voskresenskaya and
Psekups at Babinskaya
and Saratovskaya
southern elephant
Equus stenonis
Rhinoceros etrus-
cus
Cervus pliotaran-
doides
CI-1033
240
Climate and landscape
from the composition
of mammalian fauna
In Transcaucasia strong
development of steppes
on the plateaus,
resulting in migration
of southwest Asian
species to Caucasus
On Ciscaucasian plains
climate temperate and
dry; broadleaf forests
in foothills
On Ciscaucasian plains;
warm climate;
savannah and
gallery forests
243
Caspian region
Marine Lesser Caucasus upland and
beds Eastern Ciscaucasia р northern Iranian Plateau
; $ Eastern Transcaucasia
and Russian Plain
Bone-bearing sands of Adzhi-
Eilas near Yerevan
Elephas trogontherii
fallow deer
tur
Paleolithic sites on middle Zanga
and at the foot of Alagez;
fauna unknown
Bone-bearing sands at Leninakan,
upper beds
Elephas trogontherii
horse
oF Rhinoceros mercki
= giant deer
Е Camelus knoblochi
3 Bos primigenius
Bone-bearing sands of upper Marine coquina at Khurdalan
terraces of Kuma and on Apsheron Peninsula
Podkumok near Georgievsk | horse
southern elephant thinoceros
Equus stenonis
deer
Bos
СТ 1073
Sub-Khazar beds at Binagady
Elasmotherium
sibiricum
Bone-bearing sands at Leninakan,
lower beds
Mastodon arvernensis
Marine coquina at Kishla on
Apsheron Peninsula
hyena
Bone-bearing sands at Equus stenonis
= Podlesnyi and Divnyi
5 near Manych Diluvium at Tsinandali
= southern elephant Rhinoceros etruscus
= Elasmotherium AES On а
5 Я small Mustilidae
a hare
5 Rhinoceros etruscus
: Огух
Tash-Kala near Grozny Yenikent on Alazan
Elephas planifrons Trogontherium
Equus stenonis cuvieri
241
245
TABLE 63 (continued)
Black Sea region
Marine
beds Western Ciscaucasia and Г
: ы Western Transcaucasia
Russian Plain
Bone-bearing sands at
Khapry and Rostov on
Azov Sea coast
Elephas planifrons
Mastodon arvernensis
Hipparion
Climate and
Subdivision of
Quaternary
landscape from
the composition of
mammalian fauna
Warm climate;
savannahs and
gallery forests in
the southern part
of the Russian Plain
Fauna unknown
Equus stenonis
camel
CI 1056-1142
Pliocene
Upper
Eolithic
Krasnodar beds
Paleontological studies of Holocene mammals lead to a number of general
conclusions on faunal and landscape changes in the Caucasus which have
occurred since the end of the Upper Pleistoce. The absence from the Holocene
beds of those mammals which lived during the stage of maximum cooling indi-
cates that a discontinuity in ecological conditions promoted their extinction.
The paleontological materials indicate that the ecological assemblage of
the piedmont plains was the one most affected in postglacial time; mountain
fauna was probably affected to a much lesser degree.
Inasmuch as species such as cave bear, cave hyena, Panthera leo,
Equus hemionus, giant deer and others lived in the cold climate of
the Upper Pleistocene, it may be assumed that the main reasons for their
extinction were abrupt warming and increasing drought. The development
of xerothermal landscapes must have affected the fauna of the lowlands
and foothills to the east much more pronouncedly than the fauna in the
western part of the Caucasian Isthmus. A number of relevant examples of
the retreat of mesophilous species have been given under the discussion
of the Binagady locality.
The Holocene deposits, like the Pleistocene, do not contain any northern
cold-climate forms (reindeer, blue hare, arctic fox). No Holocene remains
of these species were found on the lower Don either. It becomes clear that
in the Holocene the ranges of distribution of these species moved north,
rather than closer to the Caucasus. Development of isolated pockets
inhabited by mountainous mesophilous rodents also indicates the advent of
a warm climatic phase, which resulted in shifts in the landscape zones.
Examples of such isolated pockets are: pine vole on the meadows of the
Stavropol Plateau, Transcaucasian mole vole on Mount Razvalka inPyatigor'e,
Asia Minor snow vole on Mount Kelakhan inthe dry Zuvanda valley of the
Talysh Mountains, and steppe vole on the Apsheron Peninsula. These
and other examples of disrupted ranges of distribution are reviewed in
more detail in a separate section under the discussion of the history of the
development and the present state of areas of distribution of some species.
The new developmental stage in Upper Quaternary faunal complexes is
marked by the appearance inthe Caucasus of new species of mammals not found
in the "universal" paleontological record of the bituminous Pleistocene beds
242
246
of the Apsheron Peninsula. The occurrences noted in the Holocene of jackal,
striped hyena, kulan and goitered gazelle and their present habitation of the
area* indicate that their ranges in the Caucasus are relatively young. The
southern origin and the migration of these species to the Caucasus in recent
times can be deduced from the fact that the main areas of their distribution
are to the south and east of the Caucasus, as well as from the fact that the
Species cannot survive deep and prolonged snow cover. In this species list,
jungle cat, lion and tiger (so far unknown in the Caucasus as fossils) must
probably be included. (This is discussed in more detail in Chapter III. )
Our paleontological materials indicate only one cycle of warming and
development of xerothermal landscapes, which reached its maximum at a
fairly early stage of the postglacial epoch.
It seems possible that the fossil mammals of a country as highly
diversified geomorphologically as the Caucasus may not reflect the minor
climatic changes which might have occurred within the Holocene.
The existence of a xerothermal stage was recognized by geobotanists
Maleev (1939b, 1946), Yaroshenko (1956), Grossgeim (1948) and
Takhtadzhyan (1941). According to these authors this stage was responsible
for isolated occurrences of xerophilous plants in western Transcaucasia
and in some places in the Lesser Caucasus uplands among more mesophytic
areas.
The xerothermal stage has also been mentioned by pedologist Yakovlev
(1914) and Zakharov (1935), geomorphologist Shchukin (1925), zoologist
Sviridenko (1927, 1937) and geologists Gerasimov and Markov (1939).
The changes in the landscape caused by man in the last stage of the
historical epoch are also traceable in the osteological material.
The abundance of boar, deer, elk and roe deer in the food remains in the
Sarkel beds of the Middle Ages indicates that extensive forests were
developed in the Don River valley. By the 18th century, these forests had
been exterminated and the result was the disruption of the ranges of
distribution of some European species in the Caucasus.
Occurrences of semifossil deer in the Stavropol area and the occurrences
of boar and deer in some sections of the Armenian mountain-steppe (now
arid) may to some extent provide confirmation of far fewer wooded areas
in that country in Upper Holocene time.
Studies of coals from the cultural beds of settlements of the second
millennium B.C. (Tsalka) and the 12th century A.D. (Gandzha) have proved
the existence of extensive forests in the present woodless areas of the
uplands of western Georgia near the villages of Dabi and Gomi, in southern
Georgianear Tsalka, and in Azerbaidzhan on the barren ridges of Kirovabad
(Yatsenko-Khmelevskii and Kandelaki, 1940, 1941a,b).
Our studies of the ritual collections of skulls in the mountain valleys of
North Ossetia also indicated, through the disappearance of skulls of elks
and bisons, that the forests on the sloping piedmont plains of eastern
Ciscaucasia were exterminated by the 18th century.
The paleontological materials studied provide many other data on changes
inthe ranges and dispersion of synanthropic species and on extinction of animal
game through direct extermination by man and through the indirect effects
of radical anthropogenic changes in the landscapes.
* With the exception of the extinct kulan.
243
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249
250
Part Two
ANALYSIS OF THE ORIGIN OF CAUCASIAN
QUATERNARY MAMMALS IN RELATION
TO THEIR DISTRIBUTION, ECOLOGY,
AND MORPHOGENESIS
Chapter Ш
REVIEW OF THE EARLY AND CONTEMPORARY
POPULATIONS AND RANGES OF SOME MAMMALS
The patterns of zoocenose formation can be traced in any given country
most easily by an investigation of range development, with consideration of
the ecology of separate species. Well-defined ranges form the basis of
zoogeographical division.
The existing range of a Palaearctic species may be only a relict of an
ancient range, e.g., from the Quaternary period.
Therefore, in order to reach a conclusion on species dispersal, its
ecology must be considered against a background of paleographic and
paleontologic data, current zoogeographical conditions and man's influence
during historical times.
The various factors which determine species survival or extinction
divide into those of major influence and those of lesser influence.
The investigations of the Binagady asphalt deposit (Chapter II) have shown
that peculiarities of early and recent ranges of eastern Transcaucasian
mammals depend primarily on the evolution of the Quaternary landscape.
To a lesser degree these peculiarities are also dependent on morphological
evolution and biocenotic interrelationships. It is thought that the
phytophagous mammals behave autonomously during the development and
extinction of ranges. The influence of predators on Herbivora is probably
of secondary importance to their distribution.
Theoretically the explanation of the origin and development of the range
is more easily accessible from observation of colonial species —
Carnivora, Proboscidea, Rodentia, Ungulata — remains of which are
frequently found in various deposits. Ideally, these species should be
comparatively stenotopic, thatis, associated with acertaintype of topography
having a small number of biotopes. Species thus widely distributed leave
more fossil material and more recent evidence of early ranges than
narrowly distributed species endemic to their relict ranges. Eurytopic
animals yield excellent paleontological data, particularly ifthe investigator
has a consecutive chronological series of deposits to study.
The origin of other species, the fossil remains of which are rare,
particularly the Insectivora and Chiroptera, may only be proved by means
of zoogeographical mapping, analysis of present-day ranges, ecological
analogy and extrapolations on the basis of more fully investigated specimens
of Carnivora, Rodentia and Ungulata.
An attempt has been made to trace the early and recent distribution of
both living and extinct Caucasian mammals and the factors pertaining to
their range and numbers. An investigation has also been made to determine
245
251
when these species appeared in the Caucasus and the routes taken by
immigrant species. The examples selected include various orders and
species.
Order INSECTIVORA
The paleontological age of generic series belonging to this order is quite
ancient. The Insectivora fossils are known from the Eocene. Radial
adaptation and the diversity of its relationships to the landscape are complex
in the Insectivora. The earliest aquatic and widely specialized forms
belong to 3 genera: Desmana (desman), Talpa (mole) and Crocidura
(shrew) which were found in the Lower Pliocene complex of the Stavropol
area (see Chapter II). Remains of 5 species belonging to 5 genera —
Frinaceus, teniecninus, Crecraura; ету sane ial pee
are known from the Pleistocene deposits.
The contemporary fauna of the Caucasian Isthmus includes 15 species
of Insectivora belonging to 8 genera and ecologically to three categories:
fossorial, 1 genus and 3 species (mole); amphibious, 2 genera and
2 species (desman and water shrew); andthe remainder which are more or less
specialized and associated with various biotopes (e.g., hedgehog, shrew).
Family ERINACEIDAE
European hedgehog — Erinaceus europaeus (s. lato). Fossil
remains of Erinaceus species are known in Europe from the Miocene.
Remains of the European hedgehog have been found in the Caucasus inthe Middle
Pleistocene strata of the Apsheron Peninsula and т Upper Pleistocene strata of
western Transcaucasia. The hedgehog is widely distributed today (Map 1),
found from the Caspian Sea level to an altitude of 2,500 m throughout the isthmus.
The hedgehog is absent from alpine meadows, semisteppe, semidesert, open
desert and permanent bogs апа 1$ rarely seen in upland forests; it is
most frequently observed in the lowland forests of Transcaucasia and the
sparse forests and shrubbery of the rolling Ciscaucasian plains.
This hedgehog lives in tugai thickets, gardens and oases of the semi-
deserts in eastern Ciscaucasia and Transcaucasia. The range diminished
in the Pleistocene over large areas of eastern Transcaucasia; the remaining
ranges in the Apsheron Peninsula are attributed to the postglacial dryness
of the area (according to Vereshchagin, 1949c).
The origin of European hedgehog on the Caucasian Isthmus remains
uncertain. Its range extends far to the north and south to the Caucasus. The
absence of the hedgehog from open desert, semidesert and lowland steppes
indicates its mesophilous character, its ancient associations with forest
and its probable descent from Pliocene ancestors living in mesophytic
environs of the eastern Mediterranean. Hedgehog remains from Middle
Pleistocene deposits in Transcaucasia also confirm its presence in the
Lower Quaternary, or more extactly pre-Quaternary, and the early
distribution of this species in the Caucasus.
Today the European hedgehog is spreading rapidly over the plain
concomitantly with the development of steppe and desert irrigation.
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252
Long-eared hedgehog — Hemiechinus auritus Gmel. Remains of
this species were found in Middle Pleistocene strata of the Apsheron
Peninsula, and in Upper Pleistocene deposits of the northern Caspian
lowland (Vereshchagin and Gromov, 1952). Hedgehog remains are known
in the southwest from Paleolithic layers of Palestinian caves. Bones of
the long-eared hedgehog carried into the caves by eagle owls have been
found by the author in Holocene deposits in many areas of Ciscaucasia
and Transcaucasia.
Its contemporary range includes the dry lowlands of Ciscaucasia and of
eastern Transcaucasia and the high plateaus of the south (Map 1).
In the foothills of the Greater Caucasus, the hedgehog is found inthe area
between sea level and an altitude of 300-350 m. In southern Transcaucasia
and Talysh it lives in the mountain-steppe and semidesert to altitudes of
1,600-1,800m. The animal is absent from the mountain and lower forest
zone. The largest population of the species is observed in the semidesert
zone, i.e., the Kura-Araks lowlands.
The adaptation of the long-eared hedgehog to a semidesert and steppe
environment is expressed in its food specialization: mollusks, locusts and
beetles, and, to a lesser extent, small reptiles and rodents. Low water
requirements, the ability to endure high temperatures and its use of turtle,
gerbil, suslik, fox and badger burrows associate this hedgehog with a
southern xerophilous fauna.
Extensive vertical distribution of the hedgehog in southern Transcaucasia
and the relocation of the range southward indicates the probable penetration
of the species from southwest Asia during the Upper Pliocene.
The present range of the hedgehog is increasing somewhat with the aridity
of the foothills, which has been brought about by man.
Family TALPIDAE
Мое _ lt alpa сацсавтса 'ВаЁ., i. ormenta 5 "Оба ihe
remains of the genus Talpa are known in Europe and North America
from the Miocene (Trouessart, 1898-1899a; Simpson, 1945). The earliest
remains of small moles (Talpa sp.) in the Caucasus were in Pliocene
strata near Stavropol.
Remains of the recent species T. aff. caucasica are found in
Pleistocene strata of western Transcaucasia.
In Holocene sites they are very common, especially inthe Ciscaucasian
lowlands.
Considering the early isolation of Caucasian moles from the European
mole (Talpa europaea L.) and the Mediterranean mole (Talpa coeca
Savi) (cf. map by Bobrinskii, Kuznetsov and Kuzyakin, 1944), we class them
as separate species (following Ognev, 1926a, 1928). The morphological
adaptations of both species are similar; their sympatry is shown in
Map 2. A recent study of species' ranges shows that Ciscaucasia, the
Greater Caucasus and the Lesser Caucasus are inhabited by the Caucasian
mole, with the subspecies T. caucasica ognevi Str. оп the Rachin and
Trialet ridges, whereas the eastern mole is found living in the Colchis
lowland and the Talysh area.
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254
The present distribution of the Caucasian mole is significantly related
to the country's topographical development, as can be seen from the following
features of its ranges.
In western Ciscaucasia, the animal inhabits the Don, Yeya, Chelbas,
Yegorlyk and Kuban river valleys whereas in the Kuban Plain the population
declines during arid years and survives only in more humid depressions.
There is an insular distribution of a large and varied population in the
meadows and forests of the Stavropol Plateau.
In the northern Caucasus, the mole is widely distributed in the foothills,
but inhabits only the river valleys of the Taman Peninsula and Kabarda
Plain, and is totally absent from the steppelike plains. The species is
found in oak forests of Pyatigor'e and in the Podkumok valley as far as
Georgievsk, and to the north along the Kuma River to Budennovsk,
becoming rare east of Ordzhonikidze.
The mole is common in upland forests of the northern slopes of the
Greater Caucasus from the upper reaches of the Afips Basin to the Argun,
especially along the humid Chernye Gory. It has been observed on the
southern slopes of the Greater Caucasus from Gelendzhik to Zakataly and
inhabits slopes covered by fir—maple forest in the Tsebelda, Inguri and
Tskhenis-Tskhali ravines.
In western Transcaucasia, the species inhabits the terraces of the Black
Sea coast, along riverside ridges in swampy Colchis, and to the west on
the northern slopes of the eastern Taurus Mountains.
The mole is found at medium altitudes on the Surami ridge and in South
Ossetia but not in the dry Gori depression.
In the Lesser Caucasus, the range is widespread in the zone of broadleaf
forests and subalpine meadows оп the northern slopes of the Adzhar-
Imeretian and Trialet ridges in Dzhavakhetia, and on the Bambakskii and
Bezobdalskii ridges.
Although the mole occurs in forests at altitudes of 400-1,700 min Talysh,
it is rare because of the marked aridity in summer: (In July 1945, the
Vilyazh-Chai, Vassaru-Chai and Vasharu-Chai ravines yielded only 3-4 moles
per day from 100 traps. )
It is probable that the mole lives on the northern slopes of the Elburz
Range and it survives in certain parts of the northern slope of the Sevan
Range in the upper reaches of the Dzegam and Shamkhor rivers.
Unexpectedly, it also survives in the semidesert zone of the Karayazy
Plateau east of Tiflis, from a more humid epoch (Vereshchagin, 1940a).
Here it lives in oak forests and motley-grass meadows along the banks of
the spring-fed rivulets of the Kara-Su River. (Near this area steppe vole
and red-tailed gerbils inhabit the solonchak meadows of sea lavender
and arboreal saltwort (Figure 106); this is a remarkable example of the
mosaic structure of biotopes and coexistence of hygrophilous and xerophilous
animals. )
At the upper reaches of the Akstafa-Chai and Debed-Chai, Dal! (1944a)
observed the maximum density — 42 moles per hectare — ina mixed beech —
hornbeam forest at an altitude of 1,770-1,800 m. The author observed moles
on the Karabakh Upland in the forested Terter-Chai ravine at an altitude
of 700-1,600 m, but they were relatively scarce.
The mole is not found in the dry longitudinal valleys from El'brus to
Dagestan or in the interior of Dagestan. Its absence from the eastern part
248
of the Маш Range is principally attributed to the steepness of the slopes,
the thin layer of soil covering the bedrocks of the upland and the marked
aridity in the lower part of the beech—hornbeam forest zone which results
in an irregular food supply.
The Alazan-Agrichai lowland, which is humid in winter but arid in
summer, is also characterized by an absence of mole, as are the environs
of Sevan and the Daralagez ridge.
There are no data on the presence of the mole on the Iranian Plateau and
in eastern Anatolia. Blanford (1876) did not mention moles. Radugin (1917b)
included the mole in his list of animals at the Khoi oasis (analtitude of
approximately 2,000 m), probably mistaking the mounds of the mole vole
for mole hills.
(252) Loginov (1949) obtained the following data on the relative density of the
253)
mole population from animals caught in July 1937 in various vertical zones
(Table 64).
TABLE 64. Comparative population of the mole in different
environments of the northwestern Caucasus
Number of moles
caught per day
with 100 traps
Zones
PORES Sipe OF Hoos sp descaapudoonues 40
ОЕ ЕВЕ о со оворосооюо во eens 30
Бак COMEROUS HOES 65550 нобовоооовоов 5
О ISCO со Boo moe аооов рр вос 10
FIGURE 106, Habitats of ше mole (in oak forests) and red-tailed gerbil (on meadows with sea lavender
and saltwort ) in the Karayazy forest of the Kura valley
Photograph by author, 1958
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255
This survey of the collective range of Caucasian moles indicates that
their distribution is more closely associated with a landscape of broadleaf
forests and mesophilous meadows, and less so with a landscape of motley-
grass steppes. The mole's existence depends on narrowly limited physical
properties of the soil, particularly humidity and food reserves, i.e., a
considerable biomass of invertebrates. Migrations of the mole through dry
steppe and semidesert are improbable; it is more likely that the now
separated ranges of the Greater Caucasus, Lesser Caucasus and Talysh
were formerly connected. The collective range of Caucasian moles indicates
that the areas of their distribution shared a uniform landscape only in the
Tertiary. The Caucasian mole range (most of which lies within the western
regions) was already established by the Upper Pliocene following the early
development of arid zones in eastern Ciscaucasia and Transcaucasia. The
present discontinuity of mole ranges in the Caucasian Isthmus is a result
of the ''struggle'’ between mesophytic and xerophytic environments during
continuous climatic changes and of orographic peculiarities in the Pliocene
and Pleistocene.
The mole migrations from the Russian Plain to the Caucasus and back
obviously took place in the Neogene and Upper Pleistocene through the
Trans-Kuban Plain and the Stavropol area. However, the Caucasus was
also an independent focus of mole species formation.
The ranges in Ciscaucasia, on the Armenian Highland and in the Kura
River valley near Karayazy are probably of postglacial origin. The broken
range in Talysh-Karabakh may be of an earlier, pre-Quaternary age.
The contemporary range of the mole in the Caucasus has been reduced
by forest felling and the xerophytization of the foothills and mountain regions
during grazing and plowing. Only in the western Caucasian foothills is the
direct effect of trappers on moles noticeable (Graph 1). Pelt trade does
not reflect variations of population.
Desman—Desmana moschata L. The desman bones recorded from
Pleistocene strata of southern and central Europe have been described
(Trouessart, 1898-1899a; Wolf, 1938, 1939) as subspecies of the modern
О, mosehata hungarica Kormos, D. moschata fossilis: Lartet
and D. moschata magna Owen.
In the Caucasus remains of Desmana sp. (mandibular fragment in
gravel) were foundinthe ancient alluvium of a Pliocene river in the vicinity
of Stavropol. Pleistocene remains are known from Paleolithic settlements
in the Ukraine and from Quaternary alluvium of the Dnieper, Don and
Volga rivers.
The present range of the desman includes the basins of the Don, Volga
and Ural. Only a small part of this areaalong the Don valley, from the
Manych to the Don estuary where the animal has always been rare, is today
adjacent to the Caucasian Isthmus (Map 2).
The pronounced morphological and physiological adaptations of the
desman to an amphibious existence in floodplain lakes supplied with
Invertebrates indicates its ancestral evolution in river basins with a very
stable hydrologic regime peculiar to rivers in extensive plains.
The desman probably could not exist in mountain rivers of the Pliocene
and Pleistocene Caucasus because of heavy floods and continuous erosion.
Therefore we may consider the Eastern European species as a rather early,
Upper Pliocene settler, but only in western Ciscaucasia near the Yeya and
Chelbas rivers, probable tributaries of the paleo-Don.
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256
In fact some floodplain lakes are suitable for the desman, the lower
Kuban, for instance, near the Cossack village of Grivenskaya and in the
basins of the Colchis rivers Tsiya, Tsiva and Pichera.
Order CARNIVORA
The origin and age of the range area of most Caucasian Carnivora can
be determined more or less from paleontologic data and from the
peculiarities of the range. The small number of species and the
rarity of mass destruction did not provide conditions for the formation of
natural deposits of Carnivora. However, remains of predators are found
in kitchen middens of Paleolithic man and in later settlements where these
animals were hunted for their pelts. Gromov (1948) submitted a valuable
stratigraphic analysis of the remains of Quaternary Carnivora. Fossil
Canidae, Ursidae, Hyaenidae, Mustelidae and Felidae are known from the
Cenozoic deposits of the Caucasian Isthmus. The species of these families
form the present fauna of Caucasian Carnivora.
Family CANIDAE
From Pleistocene fauna of the Caucasus 5 species of Canidae of the
genera Canis and Vulpes are known; of these two are becoming extinct.
There are 5 existing species of the Holocene fauna.
Jackal —Canis aureus Г. The remains of the jackal are often found
in fossiliferous cave deposits in southern Europe (France, Italy, Rumania),
north Africa and southwest Asia.
The difficulty of analyzing small Canidae according to their skeletons
throws any identification of small Pleistocene European dogs with
contemporary African and Asian jackals in doubt, especially since remains
of Pleistocene jackals have not been found in Eastern Europe and Soviet
Central Asia. Fossils in Acheulean and Mousterian cave deposits in
Palestine, Lebanon and Syria may be considered closely related to the
present species (Bate, 1937). In Pleistocene strata of the Caucasus, jackal
remains have not been found, indicating that the jackal appeared оп the
Isthmus recently (Vereshchagin, 1949c, 19515). Later remains of jackals
were found in early Bronze Age settlements near Anaklia, in caves near
Kutaisi and in the vicinity of the Saraibulakh ridge in Armenia (Dal', 1940b).
The contemporary jackal is mainly restricted to the plains (Map 18).
In western Ciscaucasia the jackal is rare; in eastern Ciscaucasia it
inhabits the valleys of the Terek and Sunzha from Mozdok and Grozny. On
the Black Sea coast the jackal is numerous from Gelendzhik to Batumi, but
is not found at altitudes higher than 400-500 m. The jackal is particularly
numerous in alder thickets of the lower reaches of Colchis rivers. Eastward
the range passes through the Surami which only a few animals traverse.
The largest population in eastern Transcaucasia is observed on the plains
of Zakataly-Ismailly, Khachmas and Lenkoran. In winter jackals are
numerous intugai thickets and margins of reedgrass bogs of the middle
Kura and lower Araks. In fall when jackals feed on fruit, in particular
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257
medlar, they are found at altitudes to 1,000 m in the Greater Caucasus.
They also inhabit the forest zone of Talysh to its upper edge at 1,700-
1,800 m.
In summer the jackal lives in the open semidesert while feeding on
locusts but winters there only if small hibernating bustards are plentiful.
The high arid plateaus of northern Iran and Armenia are not inhabited
by the jackal; it is also rare in the deep ravines.
Despite its apparent adaptation to a moderate climate, the jackal isa
thermophilous predator, unable to endure deep snow. Its southern crigin
becomes apparent during strong snowfalls in the Kura Lowland, when the
animal becomes helpless and unable to move or to procure food, and often
perishes from exhaustion and cold (Vereshchagin and Dyunin, 1949). The
presence of the jackal among the Holocene fauna of southern Europe and its
absence from the Russian Plain are probably explained by the difference
in the thermal and snow regimes.
The jackal is now vigorously hunted everywhere, but the population
remains large. In the 1930's, 113 jackal pelts were obtained per
1,000 km? in Azerbaidzhan, 32.4 in Georgia and 10 in Dagestan
(Vereshchagin, 1947). In the last 25 years there has been a considerable
increase in the pelt yield over that recorded in 1934 (Graph 2). In 1949 the
pelt yield increased 2.5 times because of the higher premium placed on the
jackal as a menace to cattle and an unbalancing factor in the economy. This
fact demonstrates that the jackal thrives in the existing environment of
Transcaucasia.
Wolf —Canis lupus Г. ($. lato). Small wolves belonging to
the polymorphic group С. lupus existed in the Upper Pliocene
and Lower Pleistocene of Europe. Remains of a large wolf (C. lupus
have been found in strata of the period of maximum glaciation.
Those found in Germany and Austria are especially characteristic
of the Wurm age (Freudenberg, 1914).
In the Caucasus the most ancient evidence, a mandible belonging
to a small wolf (C. tamanensis), comes from Lower Pleistocene
conglomerates of the Taman Peninsula. The relationship of this species
to contemporary C. lupus is uncertain.
Remains of large wolf appear in this locality from the Lower Pleistocene
on, and are common even now (Map 19).
South of the Caucasus, C. cf. lupus remains are found in Mousterian
layers of Palestinian caves (Bate, 1937) and in Upper Pleistocene talus of
northwestern Iran.
In the north the remains of large Middle and Upper Pleistocene wolves
are common on the banks of the Dnieper, Don, Volga and Ural and in
Paleolithic settlements of the Russian Plain and the Crimea.
The present-day distribution of the wolf on the Isthmus is general
although not uniform.
According to State figures of pelt yields, the greatest number of wolves
obtained per land-unit area is from western Ciscaucasia and eastern
Transcaucasia (Map 19). In the 1930's the yield of wolf pelts per 1,000 km?
was as follows: Azerbaidzhan — 11.6, Armenia — 9.9, northern Caucasus —
6.7, Dagestan — 6.7, Georgia — 0.8.
Most pelts delivered to the State depots came from the plains and were
taken from animals killed in winter.
1704 252
258
Тре lower reaches of the Terek, Sulak and Kuma rivers and the Kura
lowland coast of the Kyzyl-Agach Gulf are densely populated by wolves which
feed on domestic cattle, wild boar and fish.
In the alder bogs of Colchis the wolf is common in the lower reaches of
the Gagida, Okum and Pichera rivers.
In the mountain regions of the Caucasus a large wolf population can be
observed in the Caucasian, Borzhomi and Zakataly- Lagodekhi reservations.
The wolf populations settled in these areas, like those that live among the
bulrushes of the Caspian coast, make predatory expeditions uphill and
cross-country after wild ungulates. The wolf is distributed almost
uniformly in all zones of the Caucasian Reservation, but in winter it is
widely concentrated in the broadleaf forest zone. At this season large
populations of boar, roe deer anddeer abound. The zonal distribution of wolves
found in the reservation (expressed in percentages) is as follows:
alpine zone — 30%; dark coniferous forests — 32%; broadleaf forests —
38% (Teplov, 1938b). According to Teplov, the loss of young boar, deer,
roe, chamois and Caucasian buck killed in the reservation by wolves ranges
from 34-61% ina five-month period. In addition to the wild hoofed mammals,
the wolf kills hundreds of domestic animals on lands adjacent to the
reservation.
The abundance of wolves in the Borzhomi area was noted by Dinnik
(1914a). The wolf is endemic in the Zakataly Reservation as in the
Caucasian Reservation (Markov and Mlokosevich, 1935; Vereshchagin,
1938a).
In other areas of the Isthmus, especially in the east, the wolf makes
regular predations after herds of domestic animals as they graze — ш the
mountains during the summer and in the lowlands during the winter. In
Spring the greatest concentration of wolves is observed in arid foothills
where there are gulleys and rock shelters; here the young are reared.
The fluctuation of the wolf populations on the Isthmus during the Holocene
can only be represented in a general outline.
The number of wolves decreased in proportion to the decrease of wild
Ungulata on the plain but gradually increased with the development of cattle
breeding. The improvement of firearms and methods of capture and the
breeding of large dogs for herd protection have probably reduced the
number of wolves.
The sharp increase of wolf pelts after World War II when the bounty
was raised showed the stability of Caucasian populations of this animal
(Graph 3).
The Holarctic distribution of the wolf in the Quaternary, its poorly
investigated geographical variability and the imperfect paleontological data
leaves its center of origin unqualified. It is considered a Pliocene dweller
of the Caucasus.
Fox — Vulpes vulpes L. (5. lato). The Holarctic genus Vulpes
dates from the Upper Miocene. Remains of Pliocene foxes, V. meridi-
onalis Nordm. and У. moravicus Wold., attainingthe size of the present
central Russian fox, are known from Eastern Europe.
Remains of Lower Pliocene foxes have not yet been recorded from the
Odessa catacombs and the Upper Pliocene strata of the coast of the Sea of
Azov. On the Caucasus remains of У. khomenkoi with a dentition
resembling that of red and arctic foxes were found in fossiliferous strata
253
(Middle Pliocene) of the Apsheron Peninsula (Bogachev, 1938c, 1944;
Vereshchagin, 1951b). An isolated fox premolar was found in Pliocene
strata near Stavropol.
The remains of foxes of the V. vulpes group are commonly found in
various types of Pleistocene and Holocene deposits of Western and Eastern
Europe, the Crimea and the Caucasus, including Paleolithic encampment
sites.
South of the Caucasus remains of foxes have been found in Paleolithic
strata of northern Iran (Coon, 1951), Syria, Lebanon and Palestine, from
which Picard (1937) recorded V. alopex for the Mousterian and
Aurignacian, V. cf. niloticus Rupp. for the Acheulean and Mousterian,
and Canis vulpes for the Aurignacian and Mesolithic.
The present-day distribution of the fox in the Caucasus is general
(Map 20). The greatest density of the fox population is observed in the
zones of semidesert, steppe, foremontane forest steppe and highland steppe;
the density is least in alpine meadows. Foxes are rare in the mountainous
beech forests of the southern slopes of the Greater Caucasus. Dinnik (1914a)
refers to the thinned forests on the foothills and plateaus of the northern
Caucasus when discussing the fox population in Caucasian forests. In the
Caucasian Reservation the largest number of foxes, upto 43.1%, inhabits the
broadleaf forests throughout the year; a smaller number, up to 31.8%,
is found in the spruce—fir forest zone, and the smallest number, up to
25.1%, in the alpine and subalpine zone Khonyakina, 1938). This distribution
generally corresponds with the particular food supply: Muridae, birds,
insects and berries. A different situation exists in Transcaucasia.
In 1935-1936, we found indications of fox distribution on the southern
slopes of the Greater Caucasus in Azerbaidzhan in the form of fresh foot-
prints and feces. The distribution according to the vertical ecological zones
in which the foxes were found is given in Table 65.
(259)
TABLE 65. Frequency of foxes found and their tracks in different zones of eastern Transcaucasia
Number of foxes and their tracks
recorded on a 20km walk
Land types in eastern Transcaucasia
[в крабы rapa 1936
8
4
И
1
In the semidesert zone the slopes of gulleys and foothills are densely
populated in spring, whereas the seashore and foothills are populated in
winter.
The basic food in summer consists of grasshoppers, and in winter of
beetle larvae, caterpillars, steppe vole and gerbils. The seashore attracts
January
Virgin semtidesert % №. od Sy bate басс. 6
Lower forests of the Agri-Chai valley
Beech—hornbeam mountain forests
а ло в gale Old р В clone
Rocks and taluses of passes
254
259
260
the foxes because of its abundant food supply: dead birds, fish, crayfish.
The contemporary fox distribution according to biotopes is greatly affected
by man's agricultural activity (cf. Chapter VI).
The yield of fox pelts during 1930-1940 showed the densest population
of foxes inhabiting the plains of Ciscaucasia and eastern Transcaucasia.
The following numbers of pelts were obtained per 1,000 km? of tOGEILOr yi:
northern Caucasus — 276; Azerbaidzhan — 268; Armenia — 158; Dagestan —
138; (Georgia — 74.
The number of foxes is inversely proportional to the degree of afforesta-
tion and to the mountainous relief of the country (Vereshchagin, 1947). The
fox is especially rare in western Transcaucasia which is rich in forests and
mountains.
Accordingto Bakeev's records, the density of the fox population in the
Spitsevka area of the Stavropol region during the period 1938-1941 was 12.5
animals per 1,000 hectares in summer and 10.3-16.5 in winter (Chirkova,
1952).
During historical time the range and population of foxes in the Caucasus
has apparently thrived under the favorable influence of agriculture, and
even concentrated gathering has failed to reduce their numbers.
The data on pelt supply during the last 25 years show sharp variations
in the number of pelts for Ciscaucasia and fewer variations for
Transcaucasia. Transcaucasia is more diverse ecologically and topographi-
cally and therefore the fox population is more resistant. From the pelt
yields of 1925-1952, three peaks in population increase can be noted — 1927,
1936 and 1946 — with corresponding decreases in 1931, 1941 and 1951
(Graphs 4 and 5).
Considering the presence of similar factors in the Middle Pliocene of
the Caucasus and the marked polymorphism of the present species in the
Isthmus (Chapter IV), the fox should be included in the local eastern
Mediterranean species of the Upper Tertiary.
Corsac fox — Vulpes corsac L. The range of the corsac during the
Pleistocene extended over a wide steppe belt from Western Europe to North
China. Remains of this fox have been found in caves of Switzerland and
Czechoslovakia. They are known also from a series of Paleolithic
encampments in the Russian Plain and particularly from the Kodak and
Crimean caves.
On the Caucasian Isthmus corsac remains were found in large numbers
but only in the Middle Pleistocene sands of Apsheron. To the south,
particularly in Syria, Palestine and Iran, the corsac has not been observed.
Here it was replaced by other mountain-desert forms of small foxes. By
the 20th century the corsac was widespread in the Caucasus, and
particularly in the northeastern part of the Isthmus (Map 21).
The reduction of the western part of the species range probably began
in the Upper Pleistocene but has only been apparent during the last 150 years
because of the pelt trade. Since the Middle Pleistocene the corsac range on
the west coast of the Caspian Sea has receded to the north for a distance
of 700 km.
Why the range diminished in Europe and the Caucasus since prehistoric
times is stillopen to question. The data available indicate that the changes
in group structure of small steppe predators occurred in the Middle
Pleistocene. In Western Europe, for instance, a series of small wolflike
and jackal-like Canidae became extinct without leaving a trace.
255
261
At the time of the Binagady asphalt trap the corsac population was still
fairly large in the biocenosis of the semisteppe of eastern Transcaucasia,
but was subsequently depleted. The number of corsacs in relation to fox
and wolf (as calculated by the authors for the Quaternary from bone
remains and pelt yields) is presented in Table 66.
TABLE 66. Comparative numbers of Quaternary Canidae (in %)
Date and locality Wolf
Holocene, 20th century. Mean annual pelt yield
in the Northern Caucasus for the period
1935-.1940;:—= 631600) fev gat. ое 2.4
Holocene, Sarkel, 8th-13th centuries A.D. Total
number of the specimens trapped by Khazars
Banaras а cvs Бо лари Зена 15.9
Middle Pleistocene, Binagady, Total number
of specimens trapped in asphalt lakes — 327 36.7
The corsac has long been a less vital form than fox or wolf, although in
those regions most favorable for it the corsac population still exceeds those
of fox and wolf even now. For example, south of Akmolinsk in the grass—
wormwood steppe 37 corsac burrows, 3 fox burrows and 1 wolf burrow
were found in 1947; in 1948, 42 burrows were observed, 34 of which
belonged to the corsac and 8 to the fox (Chirkova, 1952, p.402).
The corsac apparently originated in the steppe region of Kazakhstan.
It is a steppe-desert animal, feeding on small rodents, birds, reptiles and
insects.
The corsac was common in the steppes of the western Manych area and
could still be found near Salsk between 1900-1910 (Dinnik, 1914a; Bogachev,
1918). At present only a few pelts are obtained from this region (Graph 6).
Steppe plowing and man's agricultural activity exerts an adverse influence
on the corsac; now the animal is common only in the region of the Terek-
Kuma sandy plains.
The range ofthe corsac, considered from both its contemporary ecology
and paleontological data, indicates that this species was a typical middle
Pleistocene migrant from the Russian Plain to the Caucasus. The migration
must have occurred during the maximum southward advance of the cold-
steppe landscape.
Family HYAENIDAE
Hyenas and Ichthytheriinae remains are known in Eurasia from the Upper
Miocene (Simpson, 1945). They are especially characteristic of
Mediterranean Pliocene deposits.
Three or four fossils and one living species of hyena are known from
the Caucasus. Earlier remains were found in a Middle Miocene deposit near
Belomechetskaya (Map 21).
256
262
Two specialized lines emerged in the Upper Miocene: Crocuta, with
carnassial teeth resembling those of large cats, and Hyaena, more
primitive and with less powerful teeth.
In the Quaternary there were two species. Their exact relationship to
the Tertiary forms is not clear.
Cave пуепа — Crocuta spelaea Goldf. This species is known from
two localities in the Caucasian Isthmus where it apparently lived during the
Pleistocene: the Middle Pleistocene strata of the Apsheron Peninsula
(Binagady) and the Upper Pleistocene deposits of the Kuban Plain (Il'skaya).
North of the Isthmus the distribution range of this hyena encompassed
enormous areas of Eurasia in the Pleistocene (Pidoplichko, 1951).
To the south the remains of this animal have been observed in the Upper
Pleistocene of northwestern Iran (Brandt, 1870; Coon, 1951).
The Middle Pleistocene Crocuta spelaea Goldf. of the Russian
Plain was considerably larger than the Transcaucasian species
(Vereshchagin, 1951b), and corresponds to the larger sizes of Herbivora
of these regions, such as bison and horse.
According to cranial features and body size, the Binagady hyena occupied
an intermediate place between the Crocuta spelaea Goldf. and the
extant C. crocuta L. of Africa. A direct relationship is quite possible.
The distribution of Crocuta spelaea Goldf. decreased rapidly in
the Upper Pleistocene, moving southwestward to Africa, but it is possible
that a modified species inhabited an area south ofthe Isthmus inthe Holocene.
Thus Crocuta crocuta L. inhabited Palestine in the Mousterian,
in the Aurignacian and in the Holocene ''Natufian'' with the reappearance
of the xerothermic climate (Bate, 1937).
This animal's extinction in Eurasia is of particular interest; undoubtedly
one factor was the gradual extinction of Proboscidea and large Ungulata in
the Upper Pleistocene. Paleolithic man could scarcely have effected the
disappearance of this hyena; even the present-day hunter with his modern
arms has not wiped out the species. Pidoplichko (1951, p.160) states that the
anthropic factor is negligible in the extinction of hyena from Europe and
Asia. The Mediterranean Sea area can be considered the native country
of Crocuta spelaea Goldf. The expansion of its range should be
investigated as well as the distribution of the mammoth fauna. The
advance of the postglacial xerothermic period was marked in the Caucasus
by the migration from the south ofa less specialized species, the striped hyena.
Striped hyena — Hyaena hyaena L. Fossils of the striped hyena
were found in Paleolithic caves of Palestine, Morocco, Algeria and
Portugal. The finds reported in France are doubtful (Wolf, 1938, p.59).
Pidoplichko (1951), following Sharff (1918), assumes that the striped hyena
arrived in Europe in the Pliocene from the southwest because land extended
from Africa to the British Isles in the Pliocene. There is no proof to date
that H. prisca Mars. de 5еггез апа Н. monspessulana Croiz. etJob.
from the Pleistocene in southern France and Н. antiqua lLankest. from
the Pliocene in England are ancestral to the contemporary species.
Remains of the striped hyena have never been found on the Russian Plain
or in Siberia. Only in Soviet Central Asia, south of Samarkand, in the
Paleolithic strata of the Aman-Kutan cave have fragments of bones and teeth
of this species been found (collection of the Zoological Institute of the
Academy of Sciences, U.S.S.R.).
The absence of striped hyena remains from Pleistocene deposits of the
Russian Plain, the Crimea and the Caucasus, especially from the Binagady
251
complex, verifies that И penetrated the Isthmus comparatively late. Striped
hyena fossils are found on the Caucasus but only in Holocene deposits of the
Apsheron Peninsula. The present-day range of a number of subspecies of
this hyena embraces Soviet Central and southwest Asia and Africa. The
range areas have a characteristically xerothermic climate. The underfur of
this hyena is only slightly developed, another indication of its southern
origin. Its ranges on the Caucasus are principally in the hot semidesert
of lowlands and intermontane valleys (Figure 107) in eastern and southern
Transcaucasia. At the close of the last century hyena was still found in
eastern Ciscaucasia, between Derbent and Makhachkala (Dinnik, 1914a;
Satunin, 1915a). Occasionally animals were observed near Vladikavkaz
(now Ordzhonikidze) (Bogdanov, 1873). In the years 1861 —1869 many hyenas
were killed near Tbilisi, Yerevan and Nakhichevan (Radde, 1899). Prior to
1930 the hyena was found in small numbers and only in the sparsely populated
areas of western Azerbaidzhan and in the western areas of Georgia
(Vereshchagin, 1942b). From 1930 to 1940 the yield was 26 hyenas; from
1940 to 1950 only 5 or 6 animals were caught. Today a small number of
263 hyenas can be found in the discontinuous desert ranges of the Kartalinia
Plateau, near Udabno and Geredzhi.
The typical habitats of the hyena in eastern Transcaucasia are the eroded
clayey areas grown withthickets of juniper and pistachio.
During the last fifty years the range and population of hyenas have decreased
veryrapidly, mainly because of a planned extermination campaign provoked by
hyena attacks on children.
FIGURE 107, Habitat of striped hyenas in the Middle Araks valley (Abrakunis)
Photograph by author, 1947
Family URSIDAE
From the Cenozoic strata of the Caucasus two representatives of Ursidae
are known: Ursavus and Ursus.
258
264
Ursavus is а Middle Miocene genus found in Belomechetskaya.
Ursus 15а lower Pliocene genus known from a Pliocene deposit near
Stavropol in the Caucasus (Maps 22 and 23).
The Quaternary fauna of the Caucasian Isthmus includes three species
of bear, genus Ursus, of which only one lives today.
Ursus (Spelaearctos) rossicus Boris. was reported in 1931
from five skeletons recovered from sandy deposits of ancient alluvial flows
(Middle Pleistocene) near Krasnodar (Figure 108).
FIGURE 108. Skull of Ursus (Spelaearctos) rossicus Boris., from Krasnodar
The remains ofUrsus (Spelaearctos) rossicus Boris. were also
observed near Kherson (Borisyak, 1931), in the basins of the Greater Irgiz
(Belyaeva, 1935, 1939a) and the Ural rivers (Vereshchagin and Gromov,
1952) and are now reported to be in caves of the northern Urals. Since
remains of this species do not occur in Paleolithic strata of the caves of
western and central Transcaucasia and in the bituminous strata of the
Apsheron Peninsula, we can assume that the southern limit of its range
was somewhere in the Ciscaucasian foothills. Ursus (Spelaearctos)
rossicus Boris. probably livedinriver valleys and steppe ravines andon
slopes of foothills and was not as closely associated with karst topography as
Ursus (Spelaearctos) spelaeus. The broad molar cusps indicate
the preeminently phytophagous nature of the animal.
We consider this bear to be a Lower Pleistocene settler in the Russian
Plain; it probably became extinct during the Upper Pleistocene.
Cave bear — Ursus (Spelaearctos) spelaeus Rosenm. During
the second half of the Pleistocene the cave bear was a species associated
with the landscape of Western Europe and north Africa.
It is known from the remains of the Il'skaya camp site near Krasnodar
in the Caucasus and from eastern Transcaucasian caves (Figure 109).
Reports on the presence of this bear in the Binagady complex by Bogachev
(1939, 1940b) and Gromova (1948) are erroneous (Map 22).
259
265
FIGURE 109. Skull of Ursus (Spelaearctos) spelaeus Rosenm. with
shortened facial part from Ablaskira cave
The slight fossilization of the remains of the cave bear from the
Vorontsovskaya cave near Khosta is very interesting. It is possible that a
small cave bear population survived the Pleistocene in northeastern
Transcaucasia.
The remains of this species are unknown in the Russian Plain, except
north of the Black Sea. Near the Samara Bend, however, a small cave
bear existed in the Pleistocene which according to the skull and dentition
was intermediate between the small and large cave bear.
On the Lesser Caucasus, in Talysh and south of the Isthmus
in Kurdistan and in Syria, cave bear remains have not been found. There-
fore western Transcaucasia must be considered for now as the southern
limit of the species range in the eastern Mediterranean.
The affinity between this species and the small cave bear should be
further investigated; there is no reason, however, to consider this Upper
Pleistocene species ancestral to the small cave bear because fossils of
both forms exist within close geological proximity. From its origin, the
large cave bear can be classified as a local eastern Mediterranean species.
European brown bear — Ursus arctos L. ($. lato). In comparison
with the cave bears the European brown bear represents a more progressive
branch which at the same time retains a number of primitive features. No
less than ten forms are recorded from the Pleistocene cave deposits of
Western Europe, north Africa and southwest Asia (Syria).
The European brown bear was widespread on the Caucasian Isthmus
during the Upper Pleistocene; this is substantiated by fossils found in the
caves of Imeretia and in Apsheron Peninsula talus.
North of the Isthmus the remains of Ursus arctos L. (s. lato) were
found in Pleistocene sands and pebbles in the river valleys of the Don, Volga
and Kama, and in the diluvium of several Paleolithic encampments on the
Russian Plain. Paleolithic U. cf. arctos and U. syriacus have been
substantiated in Lebanon, Syria and Palestine (Picard, 1937).
While remains of the European brown bear are rare in Holocene
deposits of the Caucasus, the species is nevertheless found as a regular
replacement of the large brown bear in multilayered deposits indicating its
broad dispersal during the postglaciation (Map 23).
260
266
Pictures of bears are comparatively
rare in the ancient art of the Caucasian
people. In Kobanian burials of North
Ossetia pendants and seals of bronze
representing a bear were found. A
running bear is shown on a carnelian
seal (Figure 110) from an Urartu grave
on the shore of Lake Sevan. A bear
climbing a tree is figured on a silver
container from the Maikop burial.
The contemporary range of the bear
FIGURE 110. in the Caucasus is still extensive. On
the northern slope of the Greater
: ; Caucasus it extends from Novorossiisk
from the Kobanian burials of Ossetia :
(according to Uvarova, 1900); 3 — picture in the west to the upper reaches of the
of a running bear(X 2) on a сагпеНап seal Dzhengi-Chai in the east, wherever
from Urartu graves (Orig, ) there are forests, mountain shrubs and
meadows. The greatest population of
this animal was observed in the
Caucasian Reservation. On the Stavropol
Plateau the bear was exterminated in the last century and is only rarely
observed in the forests of Pyatigor'e. There are few bears in the Kabardian
A.S.S.R. and in northern Ossetia, and none in the largest part of
central Dagestan, but they are still seen in the Deshlagar forests. The bear
was exterminated about 30 years ago in the lowland forests of Samur and
on the gently sloped Kusary Plain.
On the southern slope of the Greater Range there are now more bears
than on the northern slope. Bears are common in wooded Abkhazia and
Imeretia. There are no bears in the Colchis swamp. In Svanetia and South
Ossetia the bear is common in reaches of the Kodor, Ingur and Rion, despite
energetic attempts by the local population to exterminate the animal. In
Azerbaidzhan it is numerous in the basins of the Belokan-Chai, Katekhi-
Chai, Kakhi-Chai and other rivers, as far as the upper reaches of the
Gerdyman-Chai in the east.
In September and October when chestnuts and walnuts ripen, the bears
concentrate in forests and gardens of the Alazan-Agrichai valley. The
animals congregate on the upper forest edge when beechnuts and acorns are
abundant in fall. On the low Kartalinia, Тога and Adzhinour plateaus it lives
in the sparse forests of juniper and Turkish terebinth where it feeds on
terebinth berries, juniper and grasses, and especially on astragali. The
bears track through gulleys and along precipices. Recently bears inhabited
the tugai floodplain forests of the Kura, especially the section between
Mingechaur and the Alazan estuary.
At present the bear is uncommon on the Lesser Caucasus where it has
been nearly exterminated. It is not found in woodless Dzhavakhetia. On the
Armenian Highland it inhabits sparse juniper forests and relict oak forests
and it is rare in mountain forests of Talysh.
The greatest yield of recorded bear pelts was from eastern Trans-
caucasia. From 1930 to 1940 the average yield per 1,000 km” was as °
follows: Azerbaidzhan — 1.5, Armenia — 1.5, Georgia — 0.9, northern
Caucasus — 0.9 and Dagestan — 0.2. Most of the pelts, however, were
retained by the local population.
1-2 — bronze pendant — bear figure (1:1)
261
267
The European brown bear of the Caucasus was probably of endemic
Pliocene origin. This local origin is substantiated by Pliocene finds near
Stavropol, by its abundance in the area and by the diversity of its forms.
In different parts of the country, however, new genetic influences from the
northern and southern neighboring populations were possible. The recent
Holocene relationship of the range areas of boreal and Caucasian bears
existed in the Don valley, the steppe ravines and the scarps of western
Ciscaucasia; this may explain the resemblance of the northwest Caucasian
bear to the European bear (Smirnov, 1916a).
Man's activity, which wrought changes in the landscape, also affected
the population and range of the bear. The pelt yield increased somewhat
in 1930 then dropped and increased again in 1941 (Graph 8).
The future of the bear on the Caucasus is not promising because of
continual forest felling and hunting of bear as a trade animal and cattle-
breeding pest.
Family MUSTELIDAE
Remains of Mustelidae emerge in Eurasia in Lower Oligocene strata.
From the Pleistocene fauna of the Caucasus 6 species of mustelids are known,
one of which probably became extinct in the Holocene.
Glutton — Gulo aff. gulo Г. Glutton remains have been found in many
caves from the Upper Pleistocene of the southern and middle belts of
Western Europe. Inthe U.S.S.R. glutton remains were found in Crimean
and Ukrainian Paleolithic strata on the Kama banks and in the Altai and
Urals caves.
A fragment of glutton mandible was found in the Upper Paleolithic strata
of the Gvardzhilas cave and in the Lower and Middle Paleolithic strata of
the Kudaro I cave. This was the most southeastern habitat of the
Pleistocene species.
Smirnov (1923-1924) considered the glutton a mountain animal and
explained its extinction on the Eurasian plains by the disappearance during
postglaciation of compacted snow оп whichthis animal could overtake average -
size ungulates.
Pidoplichko (1951) explained the withdrawal of the glutton range to
northeastern Europe and its rapid extinction by ''anthropic factors.'' It is
interesting that a similar decrease in range is recorded for the arctic fox,
the reindeer and other arctic animals. In contrast, the range of the glutton
shows а wide gap between its early Mediterranean focus and its northern part,
added during the Pleistocene. The Caucasian focus of the species became
extinct only slowly because food was abundant and mountain topography and
therefore snow conditions were diverse. Many factors were responsible
for the extinction of this animal in the Caucasus, the most important reason
being decline in reproduction in the present-day phase of species evolution.
The glutton is an autochthon of the mountain margins of the Mediterra-
nean area; from this focus the animal dispersed northward and can be’
observed today on this new range.
Exactly when the glutton became extinct on the Caucasus is as yet not
determined. It is, however, possible that it became extinct there later
than in southwest Europe, perhaps in the Bronze Age.
262
Stone marten— Martes foina Erxl. The genus Martes is known
in Eurasia from the Lower Pliocene. If we include the real martens in
genus Mustela, we can assume that this collective group developed in
the Holarctic by the Upper Miocene. Mustela filholi Goudry and
Putorius sp. were reported by Bogachev (1938c) from the Caucasus in
the Pliocene deposits of the Armenian Highland.
Fragments of Martes sp. were collected by archaeologists in the
Middle and Upper Pleistocene strata of western Transcaucasian caves:
Akhshtyrskaya and Sakazhia. On the Russian Plain we found jaws of
Martes sp. inthe Pleistocene sands of the lower Kama. This species
is probably related to the pine marten. Real martens were not found in
lower streams of Russian Plain rivers, but in the Crimea they were common
in Paleolithic strata. From the Palestine Paleolithic, a M. palaeo-
syriaca Nehr. similarto М. foina has been recorded. The marten
of Urartu time from the Sevan shores was incorrectly recorded as
Тат rons ‘Sat!
The contemporary range of the stone marten is enormous: it spreads
from the shores of Spain along the middle latitudinal belt as far as northern
China and Manchuria. The Caucasian subspecies M. foina nehringi
Sat. inhabits forested foothills, mountain forests and woodless plateaus
(Map 24). On the Stavropol Plateau and on the undulating plains west of
Stavropol it inhabits shrub-grown ravines and scarp lines of Sarmathian
limestones. There is an annual yield of pelts from the middle reaches of
the Kuma inthe area of Budennovsk, andthe Greater Caucasian foothills, the
anterior valleys and central Dagestan are also inhabited by the stone marten.
It is widely distributed in the basins of the left tributaries of the Terek,
the Sunzha and the Manas, and in the Gunib District. An excellent pelt yield
was taken from the lower forests of Samur. We found this marten near
Baku, near the base of the Apsheron Peninsula.
The stone marten is absent from the fringing forests of the Kura-Araks
lowland.
In western and eastern Transcaucasia Martes foina Erxl. dwells
primarily in the foothills and lower mountain forests; it is very numerous
in the lower forest areas, e.g., in the Alazan-Agrichai valley, where
pelts are generally obtained. On the Armenian Highland and in Talysh the
stone marten is widespread wherever there are small shrub thickets and
forests. It lives in colonies in the upper mountain zone. However,
even in desert intermontane valleys, e.g., on the middle Araks, the
animal inhabits only large rocky placers and valleys, and ravines of
isolated summits such as Ilyanlu-Dag, Darry-Dag and others.
Judging from its ecological distribution the Caucasian stone marten is an
early Pliocene dweller originating in the south.
The activity of man, especially trading, affected it less than the following
species.
268 Pine marten—Martes martes Г. The remains of the pine marten
are common in Pleistocene deposits of Western European caves (in England,
France, Italy and Hungary). They are also recorded from Mousterian
strata of Palestine (Bate, 1937). It is difficult to distinguish the bones
of this marten from the bones of the stone marten which probably lives in
some Paleolithic encampments of eastern Transcaucasia.
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269
The present-day range of the Caucasian subspecies М. martes
lorenzi Ogn. is very peculiar; it mainly inhabits full-grown beech forests
on the northern and southern slopes of the Greater Caucasus, gradually
disappearing in the east. On the Lesser Caucasus it inhabits the northern
slopes of the Adzhar-Imeretia ridge, and on the Trialet ridge it is found
from Borzhomi and Bakuriani to the Belyi Klyuch. To the west, the range
extends from the northern slope of Taurus to Asia Minor (Map 25). East
of Debed-Chai and Akstafa-Chai the marten is very rare if it exists at all.
No marten pelts have been obtained from Karabakh forests during the last
25 years.
Satunin (1907b) recorded a skull of a young marten from a site located
on the eastern shore of Lake Sevan as M. martes (?). His diagnosis was
based upon delicate features and small size; this skull, which is now in the
Georgian Museum, is in fact that of a stone marten.
The range of Martes martes L. had not reached Lake Sevan by
the middle of the first millennium B.C. From the seven marten skulls taken
from the Urartu graves not one belonged to Martes martes L.
The species is absent from Talysh despite the abundance of forests and of
the fat dormouse, а very important food. Ellerman and Morrison-Scott
(1951, p.245) incorrectly claim the presence of the pine marten near
Asterabad.
The absence of the pine marten from the eastern part of the Lesser
Caucasus and the Talysh Mountains may be attributed to the late appearance
of the species in the Caucasus and the presence of an ancient woodless
barrier in the Khram basin. The competition of the stone marten in the
southeastern regions of the country should also be taken into consideration.
The greatest number of pine martens is observed in the western
Caucasus. Pelt yield per 1,000 km? for both martens* is given by pelt
supply stations for the 1930's: Georgia — 37.8, Armenia — 30.3,
Azerbaidzhan — 19.4, Dagestan — 10.3 and northern Caucasus — 7.8.
The high number of martens in Georgia is explained by the heavy
afforestation of the country. In Armenia the entire pelt yield was from
stone martens.
Donaurov (1949) observed the pine marten in the Caucasian Reservation
between 1935 and 1936. (Its summer excrements and winter snow tracks
were found distributed according to Table 67. )
Such a distribution is the result of an abundant supply of vegetable and
animal foods: during summer months — Muridae and insects; in fall and
winter — mountain ash, whortleberry, common yew and rodents.
The small population of pine marten in the lower forest is probably a
result of its replacement by the stronger stone marten. Onthe southern slopes
of the Greater Caucasus the distribution of the animalis somewhat different.
Table 68 shows the frequency of encounter of the marten activity in the
Zaketaly Reserve and adjacent lands, according to our data of 1935 — 36.
Our data and those of Donaurov for the lower and middle zones also
pertain to the stone marten.
According to Mlokosevich's observations (Markov and Mlokosevich,
1935) made over several years in the Lagodekhi and Zakataly regions, the
pine marten inhabits the upper mountain forests and is unable to compete
successfully with the stronger stone marten of the foothills. It is absent
from the woodless plains.
* The State Yield Supply recorded both marten species together.
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1270
TABLE 67. Frequency of pine marten tracks in the western Caucasus (in $)
Summer
Habitat Winter
Broadleantorests wes sisters ORES cui ae ele ое 6.9
Fir—beech forests without vacciniaceous plants 9.4
Fir—beech forests with underbrush and
VEISCHMECEOWS рай 5. Jaa a 5 tao b om ule Elole o 51.8
Upperiforestied seme и по: ее 30.9
SUbalpinekmeadows emis ks о ие
TABLE 68. Frequency of pine marten tracks on the southern slopes of the Greater Caucasus (in %)
Habitat Winter
Beech—hornbeam forest of the lower zone
Beech forests with mazzard cherry and filbert
GM (NS ООО. WOM ‘сооозонор в оо ововою
Thinned oak—beech forests of the upper zone
SUDAN SIMS WCAC Golhosdootiadcnabaouge
The pine marten is a more specialized and phylogenetically younger
Species than the stone marten as shown by its morphology and ecology.
The separation of pine marten probably occurred in Europe in the Lower
Pleistocene since it was a distinct species by the Middle Pleistocene. The
ecology of the pine marten's contemporary range confirms the relatively
late (Upper Pleistocene) migration from Europe through the Balkans and
Asia Minor.
The direct and indirect influence of man on the population and range of
Caucasian martens became visible in the last century.
Ancient Circassia produced a heavy yield of pine martens. There are
references in Dubois de Montpéreux (1843) of the yield of pelts in this area
in the 18th century and Radde (1866) records an annual yield of 500 marten
pelts from upper Svanetia in the middle of the last century. Marten-hunting
for pelt-trading was taken up in the 1860's following the settlement of the
northern Caucasus (Dinnik, 1941а; Turov, 1937). The clearing of significant
areas of forest land over the past centuries in Dagestan and Azerbaidzhan
could have brought about a reduction in the marten range. Records of the
last 25 years show a decrease in pelt yield in 1930-1931, an increase in
1937 and another in 1947. Some drop in trade in Abkhazia was observed
in the 1930's as an apparent result of the development of the fat dormouse
trade (Markov, 1938; Graph 9).
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271
Family FELIDAE
At least 10 forms of cats have been identified from Neogene deposits
of the Caucasus and adjacent areas. Machairodus sp. is known from
Sarmatian deposits near Armavir and a medium-sized Felis sp. from the
Maraginsk Upper Pleistocene site in northwestern Iran. More fragmentary
bone findings of Felis and Panthera have been reported from the
Lower and Upper Pliocene of Stavropol and the Taman Peninsula.
Six species have been recorded from Caucasian fauna of the Pleistocene:
Panthera spelaea Goldf., panther, lynx, European wildcat, spotted
cat and cheetah. And finally from the Holocene there have been nine
species identifications; of these nine, three have become extinct in historical
time.
Cave lion— Panthera spelaea Goldf. Remains of this species
of large cat have been discovered in Middle and Upper Pleistocene
Eurasian strata — from England to the New Siberian Islands (Trouessart,
1904-1905). It is a characteristic index species of the Pleistocene mammoth
fauna of the Russian Plain and of the Crimea (Gromova, 1932a; Gromov, 1948;
Pidoplichko, 1951). The southern distribution limit of P. spelaea Goldf.
was probably somewhere in the Mediterranean area. Remains of Paleolithic
cat from Lebanon, Syria and Palestine probably belong to the real lion
(Picard, 1937). There have been no findings of P. spelaea Goldf. in
Iranian caves. On the Caucasus remains of a large Pleistocene cat were
discovered in the Sakazhia cave, in loamy talus deposits in the vicinity
of Sochi and in asphalt sands of the Apsheron Peninsula (Figure 111; Map 33).
Gromov's assumption (1948) that Panthera spelaea became
extinct on the Caucasus just before the Bronze Age, which would have been
later than in other regions, is open to question. In all probability, it
became extinct simultaneously with the cave bear, woolly rhinoceros and
mammoth. Related species, such as lion and tiger, appeared on the
Caucasus as postglacial immigrants from the south.
Lion— Panthera leo Г. The subject of the lion's place in the.
Caucasian Holocene fauna and of its relationship to the Pleistocene
Panthera spelaea is very complicated. The majority of paleontologists
considered P. spelaea the direct ancestor of the contemporary lion.
Without going into details, Pidoplichko (1951) claimed that the large
Pleistocene cat was also a lion. However, asour investigations have shown
(Vereshchagin, 1951b), Panthera spelaea possessed more specialized
feline cranial features than the contemporary African lion.
A number of West European paleontologists have recorded lion remains
in caves in England, France, Spain, Germany, Italy, Poland, Sweden,
Czechoslovakia, Hungary, Morocco and Rhodesia, mostly from Holocene
strata (Wolf, 1938, 1939).
Lion remains were reported by Picard (1937) in Acheulean and
Mousterian strata in Palestine.
Gromova (1932a) mentioned teeth and metapodia of lion found in an
ancient Greek cave in Olvia; however, these remains may have come from
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212
a lionskin rug brought from the south and not, as reported, from an animal
caught on the north coast of the Black Sea.
Spendid low- and high-relief representations of lions found during the
excavations of ancient Sumer, Babylon and Nineveh verify that the lion was
very common in southwest Asia in the third to first millennia B.C. Another
series of representations of lions executedon stone by the Hittites in the
central part of Asia Minor was published by Osten (1929-1930).
FIGURE 111, Skullof Panthera spelaea Goldf., (female) from
Binagady asphalt
Lions are known on the Caucasus only from pictures and legends. Among
them are the following noteworthy findings: relief representations of two
lions on the gold plating of a beaker from the Trialet mounds, 2nd century
B.C. (Kuftin, 1941); contour images of two lions on a gold cup from a
Kirovakan burial, second millennium B.C. (Piotrovskii, 1949); contour
images of lions on rocks in Kabristan from the first (?) millennium B.C.
(Vereshchagin and Burchak-Abramovich, 1948); contour images of lions
on silver containers and sculptures on gold plates (Figure 112) found
near Maikop, second millennium B.C. (Farmakovskii, 1914); relief
representation of a lion on the gold plating of a quiver from burials on the
Taman Peninsula (Figure 113).
Representations of lions and lionesses dismembering fallow deer and
wild asses were found on a gold vial from the Solokh burial in southern
Ukraine (6th century B.C.). Mantsevich (1949) concluded that this vial was
made by Ionian craftsmen, but (despite our advice) erroneously
classified the wild asses as horses and the lionesses as leopards.
We do not exclude the possibility that the lion penetrated into Ciscaucasia
and the Russian Plain in the xerothermic period. Pogrebova (1950) formed
the reasonable opinion that most of the artistic works and representations
of animals found in the Scythian mounds were executed locally and were,
therefore, modelled on the local fauna. The find, in the strata of ancient
Tanais, of a plate of local raw limestone (43X56 cm) depicting a lion
supports this theory. This stone is kept in The Hermitage (Knipovich, 1949).
Later likenesses of lion appear on many relics of Armenian, Arabian
and Iranian culture in Armenia, Azerbaidzhan and Georgia. Lion heads
appear on the walls of the Old Derbent fortress which dates from
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273
the time of the Arab caliphate, and оп the fortress gate in Baku, which
was erected in honor of Shah Abbas in the 17th century.
Lion images are frequently found in the medieval art of the Dagestan
mountain region (Bashkirov, 1931). In Iran at the time of the Safawids,
hunting rugs were designed with lion images (Kverfel'dt, 1940).
FIGURE 112, Gold plates with figures of lions from a Maikop burial
Both the earlier and later representations of lions found on the Caucasus
could be easily accounted for by the influence of Hittite, Assyrian,
Babylonian, ancient Iranian and Arabian cultures, and even by the
importation of finished articles from southwest Asia, But a comparison
of the pictures on Kabristan rocks with Scythian articles of Ciscaucasia
suggests a different explanation.
It is possible that the distribution area of the European and of the Asian
lion was irreversibly reduced in historical time even in southwest Asia.
Herodotus and Aristotle mention the presence of lions in the Balkans
in the second half of the first millennium B.C. The lion figures in Homer's
Iliad. A Greek military division of the 5th century encountered north
African savannah animals near the source of the Euphrates (Xenophon, 1896
edit. ). These included ostriches and onagers which became extinct in the
19th century.
Surveys on the early distribution of the lion in the Near East have relied
heavily on such literary references as Sundevall (1863), Keller (1909),
Gromova (1928) and Korner (1930).
Lucian of Samosata (125 A.D.) reported a large cat (lion?!) which jumped
on the horse of a Scythian hunter (Latyshev, 1947-1948). Zhitkov refers
to the probable presence of a large cat — "а ferocious animal'' — on the
southern steppes in old Slavonic times in "Тре Lay of the Host of Igor"’
(Slovo о polku Igoreve, 1936, p.229), as does Vladimir Monomakh in
"Instructions'' (Pouchenie). However, Sementovskii (1857) understood
"ferocious animal'' to mean only a wolf. Moisei Kalankatuiskii wrote
(10th century A.D.; 1861 edit., p. 6) in what may be an imperfect
translation: "Blessed is the country of Agvan*... where are wild animals:
lions, tigers, panthers, wild asses; and a great number of birds: eagles,
falcons and their like." It is interesting that the reference to wild
asses was verified by kulan bones found in 1946 during the excavation
of a 13th-century settlement in Baku. At the time of the Arabian
caliphate and later, eastern Transcaucasia was entirely suitable for
the habitation of lions. The tugai on the Kura and Araks rivers and the
* Ancient Albania and Azerbaidzhan.
268
vast reed-grown areas оп the Mugan floodplains were densely populated by
boar and deer and herds of kulans and goitered gazelles grazed in the open
semidesert.
Herds of ungulates were attracted in spring, summer and fall to the
piedmonts grown with juniper—pistachio forests and a luxuriant grass cover.
In the hunting ode ''Shikhariya'' dedicated to Manuchekhr, the son of
Shirvan Shah, the poet Khagani (Avsaraddin), who lived in Shemakha
(1120-1194), glorifies the Shah's lion hunt, mentioning the area between
the Caspian Sea and Shemakha (History of Azerbaidzhan Literature, Vol. 1,
1943; Text of Arasla in Azerbaidzhan).
FIGURE 113. Part of gold lining of a quiver from the Semibratnoe site,
showing a lion dismembering a deer
In Farsi the word ''shir'' means lion and we can assume that the Shirvan
region derived its name not only from the ruling shahs, but also from the
lions which inhabited it.
In his poems 'Iskandarnama,'' ''The Seven Beauties'' and others,
Nizami Ganjawi (11th-12th centuries, A.D. ) constantly used the lion as a
literary allusion.
Usamah ibn Munkidh (1922-1923 edit. ) described lion hunts in Syria
and Palestine in the 12th century when lions were as common there as hares
in present-day Caucasia.
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274
At the end of the 19th century, Blanford (1876) indicated that lions had
become extinct innorthernIran, but were still living in Mesopotamia on
the western Zagros spurs and southeast of Shiraz. The Persian lion,
P. leo persica Meyer, 1826, was described from a Teheran specimen,
i.e., from northwestern Iran.
Chardin records that lions and tigers still inhabited the forests of
Mingrelia and Imeretia at the beginning of the 17th century (1735, p.51).
Chopin (1852), who lived in Armenia, stated that during his lifetime lions
were no longer observed there, but tigers were occasionally seen wandering
from the Araks River. Danford and Alston (1880) mentioned the
extermination of horses by a preying lion in 1873 or 1874 near Bilecik on
the Upper Euphrates. According to Kinnear (1920), lions were no longer
seen in Asia Minor by the end of the last century, but they still inhabited
Mesopotamia and Arabia. After large-scale deliveries of weapons to the
Near East during World War I, the number of Asian lions decreased rapidly.
In the opinion of Pocock (1930, 1939) and of Harper (1945), the rapid
extermination of the Asian lion was entirely a result of European
colonization and wanton hunting by English officers stationed in Asia, who
killed many of these animals for diversion.
Considering the above, we can assume that the lion inhabited Trans-
caucasia, penetrating there from the south in the Holocene. After a gradual
withdrawal to the southeast and southwest (Map 33), it disappeared from
this region by the end of the Middle Ages. The reduction of population and
range of the tiger and striped hyena presents similar histories. Bothofthese
animals were observed in this region until the first half of the 20th century.
Tiger — Panthera tigris Г. Apparently remains of true tigers are
known only from the caves of China and Java (Wolf, 1938). There are
indications that findings in France and Siberia probably belong to
Panthera spelaea Goldf. Tiger remains have not been found in the
central Asian Paleolithic caves of Teshik-Tash and Aman-Kutan.
The absence of tiger remains in the Binagady deposit is important for
an understanding of the history of the tiger on the Caucasus.
Today the tiger is distributed mainly in southeastern Asia. Like the
cheetah, hyena and lion, the present-day tiger is a postglacial immigrant.
The tiger spread widely in a northwesterly direction from southeastern
Asia, probably only in the Upper Holocene. It is significant that this animal
is not represented in the art of ancient Mesopotamia, but was depicted by
the Persians, although not as frequently as the lion.
Tigers occasionally appear in ancient Persian miniatures. In the 1620's
the Russian merchant Fedot Kotov colorfully described a tiger in the
Shah's menagerie in Kazvin (1852).
Reviews and critiques of the literature on the distribution of the tiger
on the Caucasus at theend of the 19th and the beginning of the 20th centuries
have been published by Dinnik (1914a), Satunin (1915a) and Ognev (1935).
In 1947 the author presented a chart of tiger distribution on the Caucasus,
showing locations where they had been captured over the previous ten years.
Eastern Transcaucasia has been the extreme limit of tiger distribution
up to the present time.
The tiger was so common near Lenkoran in the 1860's that some were
killed there each year (Radde, 1899). The tiger was frequently encountered
at the end of the 19th century on the Lenkoran Plain and in the mountain
forests of Talysh where at least one tiger was killed every year (Satunin,
270
275 1915a). At the beginning of this century the tiger population began to
decrease rapidly. In the 1920's (Figure 114) tigers were sometimes seen
in eastern Transcaucasia, although they were not inhabitants of this region.
Between 1920 and 1950 only two cases of tiger-pelt yield were recorded:
one in 1922 near Tiflis and one in 1932 in Talysh (Map 33).
FIGURE 114, Tiger in the jungles of Lenkoran in the first half of the 20th century
Soviet officers who lived in Gilan and Mazanderan in 1942-1943 told us
of repeated reports of tigers and of their own personal observations of tiger
tracks and the dismembered bodies of boars and domestic animals on the
forested northern slopes of the Elburz Range.
Deforestation, the extermination of boars (the main food of the tiger)
and tiger-hunting have created unfavorable conditions for tiger life on the
Caucasus.
Panther — Panthera pardus L. ($. lato). Remains of P. pardus
were found in the Paleolithic strata of numerous caves in Africa and India.
Remains of this species have been found in caves in Syria, Palestine and
Lebanon in strata dating from Acheuleanto Neolithic (Picard, 1937; Bate,
1937) and, in central Iran, it is recorded from Paleolithic strata of the
Bisotun cave (Coon, 1952). Wolf in 1938 reported related fossil evidence.
It is possible that a contemporary species of panther of African-European
type inhabited Western Europe during the warmer phases of the Pleistocene.
Panther remains have not been observed in the Pleistocene strata of
Eastern Europe, which in our opinion indicates that the limits of the range
were more southerly — at the Caucasus and Kopet Dag ranges — during the
Pleistocene.
A unique find of Pleistocene P. pardus remains was made in the
Paleolithic strata of Kudaro I and II caves at the head of the Rion.
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276
Тре remains of Р. pardus found in Holocene localities may have been
brought there from elsewhere, as the only other reliable reports are on
findings from Greek and Roman strata of Olvia and of ancient Theodosia
(collection of the Institute of Zoology of the Academy of Sciences in the
Ukrainian S.S.R.) and on findings in the Caucasus from the Mesolithic
of the Sosruko grotto (on the Baksan) and from the ''Dlinnokryly'"' cave
(in the Araks ravine).
The absence of P. pardus from the Middle Pleistocene fauna of
Apsheron indicates that it first appeared in central Caucasia, penetrating
the area along with the mouflon and
porcupine from the south.
During early historic time,
Р. pardus was widespread. It was
known by the Scythians on the
Ciscaucasian Plain. Bronze heads
of P. pardus have been found in the
Scythian hills of the Russian Plain
and in Ciscaucasia (Figure 115).
A golden figurine of a reclining
panther was recovered from the
Kellermess hill. A splendid small
bronze statue of a panther was recently
found in Samtavro excavations, west
of Tiflis (cf. reproduction in: Vestnik
drevnei istorii, No. 3, 1948).
At the beginning of the 20th century,
FIGURE 115, Bronze heads of P, pardus from the distribution of panther was limited
Ukraine burials by the foothills and mountain regions
of the Greater and Lesser Caucasus,
and by the lowland of eastern Trans-
caucasia (Dinnik, 1914a; Satunin, 1915c). During the last 50 years the
number of panthers has rapidly decreased.
From 1894 to 1898 eleven panthers were captured in the Kuban hunting
regions. In 1904 only two panthers were caught there, while in 1905 and
1906 none were found. The panther is rarely encountered on the Main Range
in our century. In 1912 two panthers were killed in the Zakataly region;
after that time, despite the abundance of goats, gazelles and deer, only
occasional panther tracks were seen in the ravines of the left tributaries
of the Alazan. In North Ossetia two panthers were killed at the beginning
of the 'twenties. In 1949 one panther was killed in the gully of the right tributary
of the Sunzha and in 1952 another panther was killed near Staliniri in
South Ossetia.
Panthers have been more numerous and more enduring in Talysh and
Karabakh. In the middle of the last century Radde ordered and received
twelve fresh panther pelts from the Lenkoran District over a seven-week
period. From 1930 to 1940, i.e., about 70 years later, the Azerbaidzhan
pelt-supply base received only two panther pelts and a small number of pelts
was acquired privately by individuals.
Not more than ten panthers were killed on the entire Caucasus in the
1930's, and those mainly on the Zangezur ridge and in Talysh. Regular
appearances of panther in the reed-grown thickets of the central Mugan,
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Ant
for example оп Makhmud-Chala Lake, were recorded from 1929 to 1932.
Near the village of Kharmandali, a dead female panther apparently buried
by the male was found in 1929. A male panther, which had fallen into a pit
during a snowstorm, was killed near the village of Mashtagi on the Apsheron
Peninsula on February 15, 1946 (Burchak-Abramovich and Dzhafarov, 1949).
Heavy snowfalls and storms have driven the panther away from the
mountains.
After World War II, encounters with panthers became more frequent
and pelt yield increased. Fresh panther pelts were received from the
Talysh Mountains, the Nakhichevan area and the Karabakh Upland. Thus,
in Azerbaidzhan in 1947, the yield was five pelts and in 1948, thirty-six pelts.
This population increase is an apparent result of the migration of this
animal from southwestern Iran. Present-day panther distribution on the
Caucasus is limited by the forests of the Talysh Mountains and the ridges
of Karabakh, Zangezur and the Nakhichevan area (Map 34).
FIGURE 116, Three-month-old panther from Talysh, showing
typical spottiness of the young animal
Photograph by author, 1947
The panthers which inhabited the Greater Caucasus belonged to the
pale-colored subspecies Panthera pardus ciscaucasicus, now
nearly extinct.
The Transcaucasian P. pardus tullianus Val. (Figure 116) is
brighter colored with an admixture of rust tints and shorter fur. The
panther which was killed in Apsheron belonged to this subspecies. The
rapid disappearance of panthers from the Greater Caucasus in this century
278
is undoubtedly the result of persistent hunting of these animals. Unless
panther hunting is forbidden, the animal will disappear in the next few
decades from the Caucasian Isthmus.
Lynx — Felis lynx L. (s. lato). Aspecies whichis closely related to the
subgenus F. lynx pardina, dating at least from the Middle Pliocene,
has been found in Upper Tertiary deposits of southern Europe (in Moldavia
and in catacombs in Odessa, France and Italy). A cat mandible, similar
in dentition to that of a small lynx, was found in Lower Pliocene deposits
near Stavropol.
Lynx fossils of the present-day type are characteristic of Lower and
Upper Paleolithic settlements in the forest and forest-steppe zones of
Western and Eastern Europe (Wolf, 1938, 1939; Gromova, 1948;
Pidoplichko, 1951). On the Caucasus they have been found in the Sakazhia
cave in western Transcaucasia.
There are no reports of lynx remains from the Paleolithic of the eastern
Mediterranean (Lebanon and Syria: Picard, 1937; Bate, 1937; Iran: Coon,
1952). Thus, it can be assumed that in the Pleistocene the lynx was not
distributed very far south of southwest Asia. Today the lynx range lies in
plains and mountain forests and is limited by the polar circle.
During the Holocene the lynx population was considerably reduced by
hunting, but even more by deforestation. The lynx does not inhabit the
riverain forests of the steppe zone nor the region to the south, despite the
abundance of food, e.g., hare, beaver and roe. Perhaps the flooding of
the area in the spring accounts for their absence; it may also account for
the rarity of lynx remains in deposits found along the lower reaches of the
Ural, the Volga and the Don.
The present range of the lynx F. lynx orientalis Sat. onthe
Caucasus includes the entire forested area of the Greater and Lesser
Caucasus and of the Talysh area (Dinnik, 1914a; Satunin, 1915a;
Vereshchagin, 1947d) (Map 35). In adjacent areas of Turkey and Iran
the lynx is encountered in islets of mountain forests. Generally this cat
is found in mountain regions even though small shrubs may be the only
vegetation. For instance, on the eastern limit of the Main Range many lynx
live in the shrub thickets and undergrowth near the villages of Kyzyl-Burun
and Khizy where hares arenumerous. The lynx is also seen sometimes in
woodless Dagestan and in small islands of oak forest in the Armenian
Highland, e.g., along the headwaters of the Nakhichevan-Chai near Shalbuz.
The lynx originated in the ancient Mediterranean area and today still inhabits
the nearly woodless landscape of Palestine (Bodenheimer, 1935). According
to Pidoplichko (1951), the lynx does not always inhabit forest land in
southern mountain countries.
The recent yield of lynx pelts from the Caucasus, from Ciscaucasia and
from Transcaucasia has fluctuated widely. Significant increases in
population were noted in 1928-1929, 1932-1933, 1938 and 1950 (Graph 15).
The difficulties of mountain trade curtailed lynx extermination.
European wildcat — Felis silvestris Г. Remains of small cats
(F. pygmaea Lartet, F. media Lartet, F. attica Wagner and other
species) closely related to the European wildcat are known from Miocene
and Pliocene deposits in Italy, Austria and Greece, and are particularly
associated with the Hipparion fauna. Fossils of a contemporary type
are common in Pleistocene and Holocene strata of many caves throughout
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274
РЭ
Western Europe, excluding the Scandinavian Peninsula (Trouessart, 1898-
1899a; Wolf, 1938). They are frequently found in Middle and Upper
Paleolithic strata in Crimean caves (Puzanov, 1929; Gromov, 1948).On
the Russian Plain a mandible of a small cat was recovered from Pleistocene
deposits of the lower Don (Tsimlyanskaya). On the Caucasus several bones
of this cat were found in Paleolithic strata of the Akhshtyrskaya and
Tsebeldinskoe caves. There are records of Felis silvestris L.
remains in the southeastern Mediterranean area taken from Acheulean to
Neolithic strata in caves in Palestine and Syria (Picard, 1937; Bate, 1937).
We found Holocene remains of European wildcat only in the Pyatigor'e
area. It is interesting that only remains of domesticated cats were found
in the medieval strata of the Sarkel fortress on the Don. It would appear
that the range of the Caucasian subspecies, F. silvestris caucasicus
Sat., was not associated during the Holocene with the European subspecies
in the Russian Plain.
Today the range of the European wildcat includes Western Europe, Asia
Minor and the Caucasus. It probably became extinct in the Crimea during
the Holocene. Within the Caucasus the distribution of the European wildcat
gradually declines eastwards (Map 36). There are no documented data
from the Talysh and Elburz mountains; there is, however, some ambiguous
information concerning a kind of cat seen inthe forests of Lenkoran, Gilan and
Mazanderan (Blamberg, 1853; Smirnov, 1922; Bobrinskii, Kuznetsov and
Kuzyakin, 1944) which has not been verified. Associations with forest
biotopes are quite pronounced for this species, but in Ciscaucasia wildcat
distribution is limited by the small number of surviving plots of shrubs,
forests and reeds, such as those of the Kabarda lowlands, the valley of
the middle Kuma, the Stavropol Plateau and the Kuban River delta. In the
southern part of the country, wildcat is more likely to be found in stands
of forests, while on the woodless upland of the Lesser Caucasus it is
practically nonexistent. In western Transcaucasia wildcats are numerous
in the beech forests of the Adzhar-Imeretia Range and in the alder swamps
of Colchis. Contrary to the studies made on the plain by Formozov (1946)
(and repeated by Pidoplichko in 1951), the distribution pattern of wildcat
does not show a relationship to the varying depths of snow cover on the
irregular mountain terrain.
The largest population of European wildcat is observed in the lower third
of the forest zone; the animal prefers the beech—hornbeam forests.
Caucasian Reservation data show the following distribution of 43
encounters with wildcat: 76.7% in broadleaf forests, 18.6% in dark
coniferous forests and 4.7% in the subalpine zone (Teplov, 1938a). The
principal pelt yield on the Caucasus is obtained from submontane forests.
According to paleontological data, the European wildcat is an autochthon
of the Mediterranean area and, on the basis of present distribution patterns,
can be regarded as an early Pleistocene settler from Europe which
penetrated the Caucasus via the Balkans and Asia Minor.
The hunting of European wildcats for their pelts during the last 25 years
has not affected this animal's population (Graph 16).
Jungle cat— Felis chaus Guld. The remains of this cat are known
from Pleistocene strata of India (Wolf, 1938), According to Picard
(1937), fossils have been found in Syria and in Palestine in deposits from
the Mousterian to the Neolithic. Grornov (1937, р. 86) is of the opinion that
Zi
Azilian and Tardenoisian strata of the Crimean mountain regions may well
contain jungle cat fossils. The finds in Western Europe (e.g., in Stuttgart
travertines) are insignificant. Pleistocene and Holocene remains of this
species are unknown on the Caucasus, and finds of Holocene remains
on the Russian Plain are also doubtful. In the 8-13th century strata of
the Sarkel fortress near Tsimlyanskaya two inferior humeral epiphyses
were found which were similar in size to those of the jungle wildcat.
In the 14th-century strata of a Slavic village in Poltava an inferior
humeral epiphysis of a cat resembling this species in size was reported
(Gromov, 1948), but the authenticity of this report is questionable. Our
later investigations of cat fossils indicate that Felis lybica lived in
the Crimea and on the southern Russian Plain throughout the Quaternary,
but Felis chaus didnot. It is probable thatFelis chaus only
penetrated eastern Ciscaucasia and the southern Russian Plain late in the
Quaternary (during the Holocene).
At present the main distribution area of Felis chaus is in southern
and southeastern Asia, while on the Caucasian Isthmus the range is
"undeveloped.'' This cat inhabits eastern Transcaucasia from the Caspian
Sea to an altitude of 900-1,000 m. In the west it inhabits the valleys of Kura
to Gori; it has not been observed beyond Surami. In the Araks River valley
the cat is encountered as far as the longitude of Mount Alagez. It is common
along the Caspian shore, especially near Samur and along the lower reaches
of the Sulak and Terek rivers, penetrating farther through the reeds along
the Berovskie knolls and to the Volga delta (Dobrokhotov, 1939).
In eastern Ciscaucasia this cat is found as far as Grozny in the Sunzha
River valley and as far as Mozdok in the Terek River valley (Map 37).
Felis chaus is not encountered at present in western Ciscaucasia.
The pelts obtained from the Krasnodar Territory were brought from the
southeast. In the scientific collections from this region, the jungle cat
is not represented; although the literature, beginning with Bogdanov (1873),
mentions the presence of this cat on the Kuban delta, it is yet to be
confirmed.
The data of Formozov (1946) on the habitation of this cat along the reed-
grown banks of the lower Kuban River are erroneous. The finding he cites
of a pelt and skull in the Cossack village of Grivenskaya was really of
European wildcat.
cm
FIGURE 117. Jaw of cheetah from Binagady asphalt
276
281
The greatest number of this species is observed in the lowland forests
of eastern Transcaucasia. Its favorite biotopes are reed—cattail thickets
beside lakes and marshes and tugai thickets along rivers and rivulets.
Here the cat hunts water voles and marsh birds. It is not encountered
in mountain forests, but will migrate through open places in the
Transcaucasian semidesert.
Some unknown barriers impede its penetration into western Trans-
caucasia. At the first glance it would seem that, like the jackal, the jungle
cat would prosper in the Colchis, which has a topography similar to that
of Girkan, a warm climate and many swamps. The absence of the cat from
the Colchis cannot simply be explained by the ''newness'"' of the distribution
area on the Caucasus. After all, the Surami Pass is sufficiently low to
permit passage, and predatory communities are known to settle rapidly.
The absence of jungle cat remains from the Binagady locality, the ''under-
development"' of its range on the Isthmus and, finally, the stenotopic
character of the animal and its relative adaptation to warmth suggest that
it is a postglacial settler of southern origin. The area of land on the
Azerbaidzhan Plains inhabited by the jungle cat increased with the
development of a local irrigation system and decreased with its
abandonment. Most recently, the destruction of tugai forests and of bamboo
groves and accelerated trade have caused a further decrease in range and
population. It is difficult to discover the population dynamics of the jungle
cat from figures on pelt yields because the pelts of this species and those
of European wildcat are reported together.
The granting of bounties on cats sharply increased the pelt yield. This
reflects a state of species well-being, although nutria breeding is thereby
endangered.
Cheetah — Acinonyx jubatus Schr. Cheetah remains are known
from the Upper Pliocene of southern Europe and Africa (Simpson, 1945).
The species is very rare in Quaternary deposits and until recent times
was known only from the Upper Pleistocene of China (Pei, 1939). In the
Caucasus (Figure 117) they were first revealed in Middle Pleistocene strata
of the Apsheron Peninsula (Map 38). At that time the cheetah (Figure 118)
could have hunted saigas, asses, horses and hares. Possibly its distribution
in the Pleistocene extended to eastern Ciscaucasia. The contemporary range
of the cheetah includes Africa and southern Asia as far as China. Its
preferred biotopes, according to Roosevelt, Pocock (1939, 1941) and other
naturalists andhunters, are low-lying plains and hilly savannahs, deserts
and semideserts where the animal shelters among rocks and shrubs. In
Turkmenistan and Iran it lives in the open sandy, clayey and stony desert,
but prefers river valleys grown with tugai thickets where it hunts gazelles
and hares.
There are no documentary data on the occurrence of the cheetah in the
Caucasus in historical times. It was depicted (Figure 119) on an ornament
of a silver container found in the Maikop burial (Farmakovskii, 1914). The
wearing of a collar suggests the possibility of cheetah domestication by
the time of the Scythian. On a 12th-century ceramic from the excavation
of ancient Gandzha [now Kirovabad] there are pictures of cheetah somewhat
reminiscent of those depicted by the ancient Egyptians.
The Georgian Chronicles ''Kartlis Tskhovreba" place the cheetah in
eastern Georgia (Kartlia) in the Middle Ages and, from this source, its
Diff
presence was incorporated into the local fiction. Some local authors made
extraordinary statements regarding its habitation of high mountains, e.g.,
К. Gamsakhurdia in ''The Hand of a Great Master'' (1945). Usamah ibn
Munkidh (1922-1923 edit.) mentioned that Asia Minor gazelles were hunted
282 with cheetahs in the 12th century in Syria and Palestine. In the Russian
Chronicles and also in "Тре Lay of the Host of Igor'' (Slovo о polku Igoreve)
'parduses'' are mentioned — hunting ''leopards'' (cheetahs) — which served
the Russian princes in the hunt during the 11th-12th centuries (Zhitkov,
1936). This practice was borrowed from the Persians or the Polovtsian
khans, according to Zhitkov. This is quite probable, because nomads of
the Russian Plain maintained military, cultural and commercial relations
with those of Ciscaucasia and Transcaucasia from the time of the Bronze Age.
FIGURE 118. Cheetah in the Pleistocene semidesert of eastern Transcaucasia
After Aristov (1866), Pidoplichko (1951, p.178) erroneously ascribed
these references to ''parduses'' (cheetahs) in the old Slavic records to the
leopard, an animal unsuited for this hunt. The Azerbaidzhan khans and
Armenian and Kartlian princes hunted with trained cheetahs up to the 14th
century. In 1474 Доза! Barbaro saw the hundred hunting cheetahs of an
Armenian prince.
Adam Olearius (1870 edit.) recorded that a Persian shah
hunted onagers using cheetahs in the Isfahan hunting grounds. Olearius
wrote (page 935) in connection with this: ''There* are numerous predators,
such as tigers, leopards, wolves and bears. Tigers** are so numerous
there that they are sold in herds of 10-20 animals. The tigers were
generally used for the hunt because they were rapid runners and became
so highly domesticated that the hunter could place the tiger on the horse
behind him."' Later information appears on the cheetah in Mazanderan and
throughout the Caspian forests (Filippi, 1865, Blanford, 1876).
* In the Gilan area.
** Cheetahs.
278
FIGURE 119. Representation of cheetahs оп a silver container from Ма ор burial (according to
Farmakovskii, 1914)
In Asia Minor and Arabia the cheetah was rare at the end of the last
century (Danford and Alston, 1880; Aharoni, 1930; Bodenheimer, 1935).
The cheetah probably remained in a wild state in the Kura-Araks lowland
and in the middle Araks valley until the 18th century. This is convincingly
shown by the sudden decline in cheetah range and population in present-day
Turkmenistan and its recent presence in northern Iran (southern
Azerbaidzhan).
The disappearance of the cheetah from Transcaucasia and southern
Asia is explained by the depletion of steppe ungulates and the intensive
pursuit of young cheetahs for training.
By tracing the contemporary range of the cheetah in Asia and Africa,
Harper (1945) established the reduced area of distribution and the population
decline of this peculiar cat.
Order LAGOMORPHA
lLagomorphs appeared relatively early in time. Some genera of this
order, for instance Mytonolagus and Shamolagus, are known from
the Eocene. In the Mediterranean, central Asia and Mongolia, Tertiary
lagomorphs are represented by pikas and hares (Trouessart, 1904-1905;
Simpson, 1945; Borisyak and Belyaeva, 1948).
On the Caucasian Isthmus remains of only two genera, Ochotona
and Lepus, are known from the Lower Pliocene.
Family LEPORIDAE
European hare — Lepus europaeus (s. lato). The earliest
fragmentary remains of small hares were found in Pliocene alluvium near
Stavropol and, later, in Pliocene fluvial deposits of the middle Zanga,
north of Yerevan.
279
Numerous L. europaeus remains from the Middle Pleistocene of
eastern Transcaucasia (Binagady) belong to a species closely resembling
the contemporary animal.
The Upper Pleistocene remains of European hare were found in deposits
of the lower Don. In Holocene deposits on the Caucasus, European hare
remains are often found in caves used by eagle owls, as well as in strata
of human settlements.
The contemporary European hare inhabits almost all of the Caucasian
several years (Vereshchagin, 1947): northern Caucasus — 3,300;
Azerbaidzhan — 770; Dagestan — 594; Georgia — 219; Armenia — 127.
Because most of the pelts are sold on the spot, the number of European
hare killed in Transcaucasia is actually much greater, especially in
Azerbaidzhan. The yieldonthe northern Caucasus is mainly from Krasnodar
Territory and the Dagestan plain. In Georgia almost all of the hares are
obtained from the dry eastern portion (Map 39).
The hare population of the alpine and subalpine meadows of the Greater
Caucasus is insignificant. It is more common in the dry inner valleys
of the northern slope of the range and on the steppelike alpine meadows of
inner Dagestan. Оп the Gunib Plateau the European hare rests by day in
relict pine and birch forests.
This species is not found at all in continuous mountain forests of beech,
hornbeam and fir. It is true that they penetrate the forest zone moving
upwards from the subalpine meadows along woodless slopes and ridges
and along denuded ravines, but this penetration takes place mainly as man
has changed the landscape.
The contemporary range of the European hare over broad areas of the
Caucasian Isthmus shows upon examination that these animals generally
284 inhabit dry uplands and moderately dry ravines with shrubs or sparse
forests. In discussing the biotopes of the European hare, it is necessary
to distinguish between its day resting places and its feeding grounds, whichdo
not always coincide. The natural distribution of couching and fattening
places has undergone marked modifications over the millennia wherever
man has engaged in animal husbandry.
On the Ciscaucasian Plain the European hare feeds and couches in the
open virgin steppe only in spring and early summer. If the steppes dry out
or are burned, the animals concentrate in the river valleys. In fall and
winter they feed on windblown places, i.e., on the hills and southern slopes,
and couch in ravines and shrubs.
In continuously cultivated regions and in regions of intensive grazing
the distribution of the European hare is very complicated. Almost all types
of continuously cultivated land in the Stavropol area divert the hares from
virgin land, especially when the virgin land is used for cattle grazing.
Alfalfa and grass seedlings comprise the basic diet of the hare; hares
also feed on ripened field melons, wheat and barley grains in stubble
fields — the latter a probable result of careless combine harvesting
(Kolosov and Bakeev, 1947, p.12).
In winter the hare feeds on crops where it couches. It will often couch
on fall-plowed fields, crossing the next forest belt if disturbed. Permanent
280
fattening and couching places in the forest belts are observed only where
there are very wide stands of advanced age.
The contemporary widespread distribution of the European hare in the
foothills, especially on the Trans-Kuban sloping plain, is the result of man's
activity, i.e., eradication of forests and swamp drainage.
т tern Transcaucasia, inhabiting the driest plots
of Black Sea terraces and only the cleared"forest foothill glades in humid
~Colchis. =
285
— 2 eastern Transcaucasia the maximum European hare population is
observed in the dry foothills and on the slopes of the Kartalinia, Гога and
Azdhinour plateaus living in vegetation of Christ's-thorn and beard grass.
The hares are temporarily driven out of this type of land only in winter
when large sheep herds and their dogs appear (Vereshchagin, 1942b). Many
European hares live in sparse tugai thickets, particularly in the tamarisk
zone. In the Alazan-Agrichai and Khachmas lowlands they inhabit places
where the original forests have been cut and where they can couch along
the edges of forest glades. In the flat semidesert of the Kura-Araks lowland
the European hare is very rare even insaltwort—caper communities. The
plowing of semideserts and the establishment of fields, gardens and oases
immediately promotes the appearance of the hare. It is particularly
common in vineyards and fallow fields. Alfalfa and cotton plantings also
attract the European hare (except when the cotton fields are being watered
and weeded).
On the Apsheron Peninsula the hare is found in vineyards and sand dunes
where it couches among the ephedra shrubs and sand fescue. In winter when
the hare is hunted, and when sheep are grazing in the vineyards, it
couches on plowed fallow land and in winter barley fields, which, because of
their microrelief, afford better protection inthe furrows or between lumps of
earth than winter-sown land. In vineyards the hares eat the ripe grapes
and gnaw on field watermelons when they are thirsty.
Hunting during the last decades has influenced the development of the
hare population on the Apsheron Peninsula. Individuals couch during the
day on stone fences or on ruins of houses from which they have a clear
view of approaching danger.
On piedmont plains hares couch in dry riverbeds where they hide behind
boulders in order to escape easily from jackals and foxes.
In the Talysh upland and in the Araks valley the main natural biotopes
of the European hare are bottoms and slopes of ravines grown withtamarisk,
common buckthorn and buckthorn thickets. In general, the habits of this
small southern European hare are similar to the central Asian Tolai hare.
It is characteristic of the European hare, a species of southern,
Mediterranean origin, that it reproduces the year round and maintains its
largest population in the south, its litters decreasing proportionately to the
distance it migrates northward (Vereshchagin, 1938b, 1942b; Kolosov,
1941; Kolosov and Bakseev, 1947).
The hare population is greatly affected by tularemia, epizootics,
coccidiosis and, more rarely, early frosts (Kolosov and Bakeev, 1947).
These population fluctuations affect the trade to some extent (Graphs 17-18).
The prime cause of the decline in hare population in recent years is year-
round poaching at night with automobiles.
The cultivated terrain of the Caucasian Isthmus will promote an even
more valuable trade in European hare in the future.
281
286
Order RODENTIA
More than 40 species of rodents are actually known from Cenozoic
deposits of the Caucasian Isthmus and the southern Russian Plain. Holocene
fauna of the same areas account for no less than 50 species identified as
belonging to the families Sciuridae, Castoridae, Myoxidae, Dipodidae,
Spalacidae, Muridae, Cricetidae and Hystricidae.
The features of the ranges and the representatives are examined below.
Family SCIURIDAE
Only four species of Sciuridae are known from the Quaternary fauna of
the Caucasus. Of these, the marmot is extinct; two susliks and the squirrel
still survive.
Marmot —Marmota sp. Accordingto the finds made inthe KudaroI cave
in 1957 ата of marmot inhabited the Greater Caucasus inthe Pleistocene and
Holocene. Marmot bones were found in Upper Pleistocene deposits in the
basin of the Khram River in the Lesser Caucasus near the village of
Orozman. Some relatively ''fresh'' marmot skulls resembling those of the
steppe marmot were obtained by geologists from the karst funnel in the
Belaya river ravine (Map 40). It is possible that the Caucasian marmot
became extinct in the northwestern Caucasus at the beginning of this century
(Vereshchagin, Geptner, Stroganova, 1959). The find of a small suslik,
Marmota (?)sp., in lower Quaternary strata near Akhalkalaki is
noteworthy.
Little suslik —Citellus pygmaeus Pall. ($. lato). The origin and
distribution of the suslik in the Caucasus have been studied by faunists,
zoogeographers and parasitologists but remain unclear. More paleonto-
logical, morphological, zoogeographical and ecological data are required
for the task.
The genus Citellus is known from the Miocene and Pliocene of North
America and from the Pliocene of Eurasia (Simpson, 1945).
Remains of marmots of C. pygmaeus type were determined by
Buchner and also by Pidoplichko (1951) from Quaternary deposits at Nogaisk
in the Zaporozhe Region.
Upper Pleistocene remains of Citellus pygmaeus have been found
only within the limits of the present-day range on the Eastern European
plain. Vinogradov and Gromov (1952) distinguished an earlier form —
С. pygmaeus musicoides — апа amore recent form — С. pygma-
eus caspicus. The identification points up the probable retention of some
archaic features of the mountain subspecies.
From Lower Holocene, Mesolithic strata, suslik remains are known from
the first Baksan gorge. More recent bones of little susliks from eagle
owl pellets were collected by the author on the outskirts of the Caucasian
Range: in the upper reaches of the Kalaus, in the Stavropol area and in the
vicinity of Makhachkala.
The contemporary range of the little suslik includes Eastern Europe,
the northern Crimea, the Ciscaucasian plains and the plains of northern
central Asia within the steppe, semidesert and desert zones.
282
There are reasons to consider the Neogene of these areas аз the origin
of this species without resorting to Pidoplichko's hypothesis (1951) of an
earlier migration from the southern Caucasus.
In the 1950's, the southwestern limit of the little suslik on the Cis-
caucasian Plain crossed the Manych depression in the lower reaches of the
Yegorlyk and, following the Stavropol upland from the north, penetrated
the upper reaches of the Kalaus far to the southeast near the village of
Sultanskoe, 15-18 km from the Kursavka station. Farther eastward the
southern colonies extended to the Kuma near Vorontsovo-Aleksandrovskoe
and continued parallel to the latitudinal course of the Terek. From the
Terek estuary to the north, the range of this rodent continued unbroken.
An isolated colony is located between the lower reaches of the Terek and
the Sulak.
On the right bank of the Sulak the suslik is dispersed over the foothills
from Chir-Yurt to Buinaksk, while on the maritime terraces on the Caspian
it penetrated southward nearly to Kayakent.
Suslik populations exist in more isolation from the main basin of the
range in the latitudinal valley between the Sunzha and Terek ranges, west
of Grozny, and deeper in the ravines of the upper reaches of the Kuban,
Malka, Baksan, Chegem and Bezingi Cherek rivers (Map 41).
The territory between the Sunzha and Terek rivers inhabited by the suslik
in the late 1930's comprised four sections with a total area of approximately
1,000 ha (Ognev, 1947).
The little mountain-dwelling suslik was recorded by Ménétries (1832)
who noted the animal on the northern slope of El'brus; the range was traced
and described by at least 14 zoologists from the end of the last century to
the first half of the present one.
According to Sviridenko (1927, 1937) the westernmost colonies of the
mountain suslik were disposed along the Uchkulan ravine and itstributaries.
The uppermost somehow did not reachthe estuary of Makhar and Dzhalp- Kol,
and down the ravines they spread no farther than the Khudes estuary.
287 The largest habitat was and is in the upper reaches of the Malka and its
tributaries: Khasaut, Garbazi and Bolshoi Taluko. This is a vast plateau,
gently sloping north andeast from altitudes of 1,500-2,500m. Sviridenko (1937)
followed the junction of this range with that of the right bank of the Malka
at an altitude of 3,072 m through the Buruntash Pass where the animals live
not far from the El'brus glaciers. From here the susliks probably entered
the Uchkulan basin.
In the 1930's the suslik spread along the Khasaut to Narzan, a natural
boundary, and to the Taluko Basin and the southwestern slope of Mount
Kinzhal but not down into the Malka valley.
In the east the suslik colonies are at present dispersed along the ravines
of the Kurtyk and the Dzhovurgen, which are the left tributaries of the
Baksan. In the Baksan ravine, the suslik is found 1.5-2 km above the gorge
cut by the river in the Skalistyi ridge. From here, the suslik steadily
spread in 1952 along both slopes of the ravine as far as the village of Nizhnii
Baksan (Figure 120), and is especially numerous near the village of Byllym.
Above Nizhnii Baksan there is another valley colonized by susliks which
is separated by 8 km from the nearest colony on the left slope of the ravine.
The animal's penetration into the Verkhnii Baksan ravine was probably
283
accomplished by way of the Kurtyk River ravine, which is the left tributary
of the Baksan from the west. In the upper trough of the Baksan the species
was observed in 1952 in the Tegenekli area and even somewhat higher,
although Sviridenko (1937) did not find suslik even at the Adyly-Su estuary.
FIGURE 120. Habitat of the little suslik in the Baksan ravine
Photograph by author, 1952
Through the valley of the Dzhigiom River, the right tributary of the
Baksan, the suslik penetrated the Chegem valley in which it lives at a very
288 high altitude especially on the Bashi-Buzu-Su tributary. From the northern
slopes of Mount Chegem-Bashi it spread into the Bezingi Cherek
ravine.
In the Balkar Cherek, the suslik has not been encountered, although there
are places suitable for it in this area, for instance, near the villages of
Kunim and Zilga.
The suslik has never been found in the Urukh ravine. In 1947-1948, the
author travelled through the entire Urukh area, including its tributaries,
through the Balkar Cherek and the Cherek Rion pass without finding any
suslik, and local inhabitants reported that suslik had never been seen in
their area. Dinnik did not find them even in 1883 (1884a, 1884c). Thus the
Rossikov report (1887, p.44) of encountering a mass of susliks in
mountainous Digora is probably purely imaginary. Ognev, citing Rossikov
(1947, p.139), added confusion by placing Mount Kun'-Ityg and the
Zhelanoko ridge (i.e. , Dzhinal) in North Ossetia, whereas they are located
in the Malka Basin (i.e., the Kabarda piedmont).
The present geographic variations of the little suslik range within the
Caucasian Isthmus (Chapter 4) does not account for the isolation of separate
1704 284
289
parts ofthe early range of this animal оп the Caucasus, because we simply
do not know how rapidly the formation of separate features of the species
in different environmental conditions takes place.
A study of suslik distribution throughout its total range clearly shows
the early adaptations of the species to artemisia semidesert. It is precisely
in this zone of artemisia semidesert, clayey soils and specific plant
groupings that the greatest suslik population density is observed.
Considering the Neogene origin of the artemisia semidesert landscape
on the northern margins of central Asia (Lavrenko, 1938) and the mountain
xerophytes of the eastern Mediterranean (Bush, 1935; Grossgeim, 1948),
we may assume that the age of the indicated adaptations is at least Upper
Pliocene.
Suslik penetration into the true artemisia steppe is only spotty. Birulya
(1941) assumes that the factors limiting its distribution are the high, closed
stands of homogeneous grass, the heavy spring thaws and the stagnant,
overheated air in the feathergrasses.
He further stated — and correctly so — that during the course of evolution
the animal adapted to open biotopes. On the level steppe with a low grass
stand, the colony could maintain prompt signalling of mutual danger, a
complex system of burrows and normal heat exchange conditioned by moving
ground air. It is significant that among mountain susliks of the Caucasus
these early adaptations have not disappeared but have only changed somewhat.
Susliks settling in the mountains on the moraine or on alluvial gravel
have adapted to burrows dug under large boulders, thereby preventing the
collapse of the burrows in the sandy ground and utilizing the friable spaces
between cobblestones.
On mountain-steppe meadows the suslik generally lives in colonies in
open places near rock outcrops and digs burrows through the vertical walls
of paths worn by cattle grazing on the slopes. This habit of digging burrows
in different kinds of vertical walls is also characteristic of the Volga-Don
steppe suslik.
The arid floors of the Baksan and Chegem ravines have environmental
conditions resembling those of the desert-plain and for this reason are
more constantly and densely populated by susliks. The immigrating
population decreases rapidly in proportion to the altitude. Thus, on Baksan
near Byllym (August 1952) we obtained the following data on suslik population
by altitudinal zones (Table 69).
TABLE 69, Density of the suslik population in the Baksan ravine
Artemisia-thymus Ry:
Festuca-artemisia
grouping on the upper,
third terrace on stone
placers; altitude
1,200 m
Subalpine meadow with
motley-grass, in some
places with outcrops;
southem exposure of
slope approximately 30°;
altitude 1,850 m
grouping on the lower first
terrace and in places
where there are small
stone heaps, stone fences
and weeds; altitude
1,100m
Number of burrows per
285
290
Sviridenko's data (1937) оп the burrowing habits of the suslik can be explained
only by his tendency to emphasize the morpholgical-ecological isolation of
the mountain suslik from the plain suslik. Thus, he claims that susliks
will tolerate arboreal habitats, burrowing deeply in forest glades, but
digging shallower burrows in sun-warmed artemisia-grown ground.
The ages and causes of isolation of suslik colonies in Ciscaucasia are
various. The isolation of the extreme southeast colonies in the Dagestan
piedmont is caused by the more or less constant flow of the lower courses
of the Terek and Sulak rivers. Their frequent bifurcation and meandering
create fairly wide land areas, grown over with tugai forests, and reed-grown
lakes which are ecological barriers for susliks. When а river changes
course, the habitat of the animal population previously located on one bank
is left on the other bank. It has also been experimentally proved that susliks
can swim across narrow rivers (to 150 m wide) and can cross them on
bridges (Sviridenko, 1927, 1937). Isolated instances are known of gravid
susliks being transported alive over considerable distances by birds of prey,
suchas Larus argentatus.
For various reasons, the isolation of suslik colonies located between the
Sunzha and Terek ranges seems to us to be quite ancient. In the first place,
the susliks from the Alkhan-Churt valley are morphologically different
from those of the Kuma and of Dagestan — they have been reported as
subspecies C. pygmaeus boehmii Krass. Secondly, it is known that
the Terek cut through the Terek gorge between Zmiisk and Terek at a period
no later than the Middle Pleistocene. From the Malka estuary to the Sunzha
estuary, the Terek is a powerful river with a strong and straight current
and nearly devoid of oxbow lakes. Well-developed forest belts of old
galleries unsuitable for suslik settlement are disposed along the Terek, and
the northern slope of the Terek Range is covered by forests impassable to
susliks.
The eastern approach of susliks to the Alkhan-Churt valley was barred
byawide strip of broadleaf and tugai forests on the Gudermes sloping plain
and by the Argun, Gudermes and Aksai rivers.
Therefore the formation of the Terek-Sunzha colonies should be
correlated with the formation of the festuca—artemisia and andropogon—
artemisia steppe landscape of the Alkhan-Churt valley, i.e, at least with
the Middle Pleistocene.
The origin of the Elburz mountain colony of susliks is more enigmatic.
Up to the present, Shchukin's hypothesis (1925) has served to a greater
or lesser extent as the basis for various modifying theories on the
penetration of suslik into the mountains. Shchukin differed his hypothesis in
answer to statements of Kuznetzov (1890) and Krasnov (1893-1894) on the
penetration of dry mountain vegetation along the interior, longitudinal
valleys of the northern Caucasus to Dagestan. He implied the penetration
of suslik from the plain along the transverse ravines after the withdrawal
of the glacier on the periglacial steppe, and a subsequent cutting-off of the
colony from the north by the establishment of a forest strip. However, the
question of the origin of the mountain suslik cannot be resolved so simply,
because in Elburz the animal inhabited not only the valley floors, but also
the passes of meridional ranges. Besides which, the colonies disposed
along the low ridges of the Malka Basin were not cut off from the north by
a forest strip; the foothills of this region have always been woodless.
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Still Sviridenko (1927) assumed that, аз ап ''emigrant from central Asia,''
the little suslik could have appeared on the steppes of the Volga-Don and
in Ciscaucasia only after Khvalynsk time. Even from our viewpoint, the
fairly recent establishment of the suslik range in Dagestan is confirmed
by the absence of the species from central Dagestan where living conditions
are very favorable for it. Later, after studying the peculiarities of suslik
distribution in the mountains, Sviridenko (1937) concluded that they appeared
in the mountains of central Caucasia during the time of the third Pleistocene
maximum glaciation (Riss) — i.e., long before the Khvalynsk transgression —
thus ''broadening the open-steppe ranges.'' Sviridenko's conclusions
were based on Tutkovskii's rejected hypothesis (1909) of an arid climate
in preglacial time. Contrary to this view, there is no place on the Caucasus
where glaciers and foehns have not formed periglacial steppe, and
contemporary glaciers are often embedded directly into pine and mixed
forests (e.g., in Karaugom and Tseya). The formation of a dry-steppe
zone within a mountain system is associated with fringing, isolated ridges
which act as rain screens, rather than with any glacial influence. For
example, central Dagestan is very arid, but has no glaciers. What is of
importance is Sviridenko's assumption of the survival of mountain susliks
from the time of the major glaciation of the Caucasus, which contradicts
earlier geologic and geomorphologic datatothe effect that enormous glaciers
covered places inhabited by susliks. Also of significance is Sviridenko's
zoogeographical structure which coincides well with Bush's concept (1935)
that the mountain-steppe vegetation of the longitudinal valleys ofthe northern
Caucasus derives from the ancient mountain xerophilous vegetation of the
entire Mediterranean area.
Ioff (1936), who accepted Shchukin's hypothesis of the postglaciai
penetration of the suslik into the mountains, was inclined to explain the
entire history of discontinuities in the Ciscaucasian range of the suslik by
extreme fluctuations in population, colonization and extermination over
great expanses.
In point of fact, in the early ‘fifties, the break in range between
the southern suslik colonies in the upper reaches of the Kalaus
and the Bermamyt slopes was only 92-95 km on a straight line and that
between the Kuma Steppe and the Baksan ravine colonies was only 120 km.
This break may mend because the distribution of susliks to the south has
accelerated in the last decades under the influences of human activity and
the naturally arid climatic cycle. Previously it was thought that the mass
settling of susliks on the Caucasian steppes began, as it did in the former
Astrakhan Province, about 1900-1901 (Pirkovskii, 1913; Sviridenko, 1927).
However the suslik probably inhabited this area earlier, but was not
reported because the country was so little cultivated and, mainly, because
its presence was not investigated. There could also have been local
population variations as reported by Lebedev (1912) and Satunin (1920, p. 81).
All of this, however, neither excludes nor contradicts the established fact
of the suslik's rapid advance southward in present time (Sviridenko, 1927;
Romanova, 1936; Babenyshev, Birulya, Besedin, Golosovskaya, Egorov,
Korf, Yanushko, 1937).
In 1952, the author found susliks near Sultanskoe and Mironov. In the
same year, Pavlov, Pushnitsa and Shiranovich together published a chart
which indicates a shift of suslik range southward along the entire border
for some kilometers.
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292
The settlement is advancing southward at a rate of 2.5-3 km рег year.
At this rate, the plain susliks may reach the mountain colonies of the upper
reaches of the Baksan and the Malka within 30-40 years!
From geological data (Vardanyants, 1939, 1948; Nikolaev, 1941) we
can assume that the suslik appeared on the Greater Caucasus at the
beginning of the Quaternary when the recession of the Apsheron sea and
a considerable peneplanation of the ranges created areas of the Caucasian
Isthmus suitable for wide distribution of steppe and desert species of flora
and fauna. By concentrating in those parts of longitudinal valleys and
southern cuesta scarps with mountain-xerophilous vegetation, the suslik
could have survived the maximum glaciation in the mountains, although
the possibility of Pleistocene and Holocene migrations cannot be excluded.
The later settlement of the suslik was dependent upon the animal's
surmounting the ecological barriers which the forest and meadow zones in
the Chernye Gory mountains and on the Pastbishchnoi and Skalistyi ridges
presented. Suslik penetration into the mountains did not, in our view, follow
the transverse ravines of the central Caucasian rivers, as proposed by
Shchukin (1925). The ravines of the Skalistyi ridge east of Chegem have
always been narrow and humid, and impassable to the suslik for distances
of 8-10 km. These gorges became deeper with the tectonic cycles of the
Pleistocene and attained maximum development in the postglaciation.
Perhaps it is because of this barrier that the suslik is not found in the
xerophytic zone of the upper reaches of the Balkar Cherek, Urukh, Ardon,
Fiagdon, Gizel'don, Sunzha and Argun, nor in the whole of central Dagestan,
although the living conditions are certainly no worse than those below
El'brus.
The suslik's presence in the Alkhan-Churt valley as contrasted with its
absence in the valleys of the right tributaries of the Sunzha is especially
significant.
The suslik's relatively recent appearance in the mountains is indirectly
indicated by the feeble development of the eastern range along the
longitudinal valleys. The Balkar Cherek ravine is an example of an area
which the suslik has not had sufficient time to populate.
During the time of suslik migration from the plain to El'brus, the path
of distribution probably lay east and west of the forested Pyatigor'e area,
which was a barrier of sorts to animal penetration southward. It is likely
that suslik bones deposited by eagle could be found in caves in this area.
In 1952 we made a paleontological field trip into the Pyatigor'e area
and found that no suslik bones had accumulated over the past 1,500-2,000
years in the regions of Zheleznovodsk, Pyatigorsk and Kislovodsk. This,
however, does not mean that there were no susliks in the Pyatigor'e area
in the postglacial xerothermic epoch or in the Pleistocene, particularly
since suslik remains have been discovered in Lower Holocene strata in the
Baksan gorge.
The question of the origin of susliks in the Caucasian mountains could
possibly be clarified further by more intensive investigations of shelters
and caves in the mountains of Razvalkaand Verblyud, andthe gorges of Kich-
Malka, Kuma and Podkumok; the acclimatization of mountain susliks on
the plain and of plain susliks in the mountains should also be investigated.
During this study it should be possible to trace the rate of ecological and
morphological evolution for both forms. For the present, the bestestimate,
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based upon acknowledged facts, for the settlement in the mountains of the
suslik from the Ciscaucasian plains is the relatively Upper Holocene.
In the Recent epoch, man, by his activity, has unknowingly contributed
to the rapid settlement of the suslik in the mountains, as well as on the
plains (see Chapter VI). Efforts at extermination and the pelt trade (Graph 20)
were not enough to impede this process in its early stages.
Modern extermination methods and accelerated cultivation of the steppes,
the development of shelterbelts and of artificial irrigation systems will
virtually eliminate the suslik from the Ciscaucasian plains.
Asia Minor suslik — Citellus citellus L. Fossils of suslik
C.citelloides Kormos have been reported from Pleistocene strata in
European caves (Wolf, 1939), and Citellus sp. from the Upper Paleolithic
of Syria (Wolf, 1939). In the Caucasus, only more recent remains of
C. citellus L. are known from Holocene alluvial loams in the Zanga
ravine northwest of Yerevan (Dal', 1949a).
The contemporary distribution of the southeastern subspecies
C. citellus xanthoprymnus Benn. includes the Armenian Highland
and Asia Minor.
In Armenia the suslik occupies a large area of the Araks valley, from
the region of the Kara-Burun station and Alagez to the village of ПИ,
northwest of Leninakan; the upper valley and the Pambak River Basin to
the Amamla station; and the western and northern slopes of Alagez (Map 41).
The altitudinal distribution of the animal occupies a zone approximately
1.5 km wide within an altitudinal range of 1,100 to 2,700 m (Dal', 1948b;
Avetisyan, 1950). In the mountains, it lives in meadow-steppe, or even
in subalpine meadows. In the Araks valley it is found in the mountain-
artemisia semidesert, but not in the desert valley of the Araks.
In view of the considerable ecological variability of the Asia Minor suslik
(Avetisyan, 1950) it should be considered endemic to the submontane and
mountain landscape of arid and relatively hot regions. The range area
includes Asia Minor, the Aegean and the Balkans. The former habitat of
the suslik in the Zanga ravine indicates a wider distribution during isolated
epochs of the Quaternary. The present-day distribution in some areas
suitable for the habitation of Asia Minor suslik was achieved only
circuitously; for instance the arid Tertiary foothills bounded on the north
by the Kura lowland were reached by way of land lying east of the Karabakh
Mountains. The main causes of the feeble development of the Caucasian
range of this species, in comparison with that of the steppe vole and the
Asia Minor hamster, are its more conservative living habits and its lesser
migratory ability.
The local breaks and contractions in the range of this suslik on the
Armenian Highland may be a result of orogenic processes and lava flows
which covered large areas. At the time of cooling and glaciation, the upper
border dropped at least 400-500 m. It is possible that the suslik also
inhabited southern Dzhavakhetia in the Pleistocene.
In the recent epoch the distribution of the Asia Minor suslik expanded
in the upper zone and shrank in the lower as a result of pelt trading and
chemical extermination (Avetisyan, 1949).
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Family CASTORIDAE
Castoridae remains are known from the Lower Oligocene of Eurasia and
North America. In their adaptations the beavers greatly resemble the
terrestrial porcupines and, to a lesser extent, the amphibious voles and
Octodontidae. Miocene-Pliocene fossils of the genera Steneofiber,
Amblycastor, Trogontherium and Pliocene-Pleistocene fossils
of the genus Castor are numerous along a wide belt of the northern
Mediterranean and central Asia stretching from the British Isles to Mongolia.
Fossils of four beaver genera, Steneofiber, Amblycastor,
Trogontherium and Castor (Map 42) have been found on the Caucasian
Isthmus.
European beaver —Castor fiber Г. Remains of European beaver
were first found on the Caucasus in Upper Pliocene conglomerates of the
Taman Peninsula. The Taman beaver, C. tamanensis N. Ver., from
the Sinyaya gulley site, a natural boundary, was probably the direct ancestor
of the contemporary European beaver (Vereshchagin, 1957a). Other
Pliocene representatives of the family Castoridae —Steneofiber,
Amblycastor and Trogontherium —also known from the Caucasian
Isthmus, do not have direct affinities with European beavers.
A European beaver of a completely contemporary type lived in the
Caucasus during the Upper Pleistocene and the Holocene. Its remains are
found in the caves of Sakazhia (Gromov, 1948) and Uvarova in the vicinity
of Kutaisi, in the Neolithic strata of the Sagvardzhile cave, and in Upper
Bronze Age strata in the Samtavro burial fields in the vicinity of Mtskheti. *
Thus the beaver habitat in the Holocene on the Rion, Kvirila, Kura and
Aragva tributaries has been documented (Figure 121):
FIGURE 121, Remains of European beaver
1 — Femur from Samtavro deposits (first millennium B,C, ); 2 — pelvic
fragment from the Neolithic strata of Sagvardzhile cave; 3 — pelvic frayment
from the Paleolithic strata of the Uvarova cave
o
Information on beaver fossils from Sagvardzhile and Samtavro was kindly provided by М.О. Burchak-
Abramovich, Doctor of Biological Sciences,
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The remains of European beaver are very common in Quaternary deposits
on the Russian Plain, but in the Crimea they have been found only in strata
of the Upper Scythian epoch. South of the Caucasus, there have been no
validated findings of beaver remains; Bate (1937) reports none from caves
in Palestine, Syria and Lebanon.
Beaver fossil identifications from Iranian caves (Coon, 1951) are doubtful
because these fossils were probably mixed with porcupine remains.
The great number of recent beaver remains in the lower Don floodplain
(Sarkel near Tsimlyanskaya, Novocherkassk) indicates that the contemporary
habitat of this animal оп the rivers of the Cis- and Trans-Kuban plains is
quite probable. Until the 18th century, perfectly suitable beaver habitats
were located on the Psekups and on several small left tributaries of the
Kuban and even of the Terek.
Published data on the presence of beaver in the Caucasus during the
present epoch are numerous and well-known.
The beavers of Scythia and Colchis were often mentioned by ancient
authors (Herodotus, Strabo). More recent references to beavers
in Mingrelia and generally throughout Georgia can be found in Lamberti
(1654) and inChardin (1686), in the Georgian Chronicles ''Kartlis
Tskhovreba, '' and in the ''Geography of Georgia'' by Prince Vakhushti
(1904), written at the beginning of the 18th century. At the end of
the 18th and the beginning of the 19th century beaver pelts and
"castoreum'' were probably important to the import-export economy of
Georgia. This is apparent from the status of Tiflis as atrade center in 1803.
Giildenstaedt (1879) and Pallas (1831) recorded the probability of beaver
habitation on the Kuban and the Sunzha. Ménétries sent a beaver skull, found
on the bank of the Sunzha, to Academician Brandt for his collection. Rovinskii
(1809) mentioned that beavers were occasionally observed onthe Kuban.
Nordmann (1840) recorded several beavers killed in the 1730's in the Notanebi
Basin and in Colchis, and indicated the beaver's presence on the Terek.
Hohenacker (1837) mentioned the Araks valley as a habitat of beaver.
Chopin's data (1852, p.807) on beavers and ''castoreum' obtained in
Armenia, Turkey and Iran are questionable.
All these data are to be found in Brandt's monograph (1855).
The beaver skull preserved in the Zoological Institute of the Academy of
Sciences U.S.S.R. under No. 6330 (''Ménétries, 1831"), represented by
Brandt (1855), has no trace of water sluice, river sand or silt, and has been
thoroughly prepared and scraped with a knife. Perhaps this animal was
killed not long before the arrival of Ménétries in Sunzha, or was gathered
by him from a northern location en route to or from the Caucasus.
Ménétries himself did not mention (1832) the beaver among the Caucasian
animals (! ).
Radde (1866) validated beaver habitats in Upper Svanetia, in the Tskhenis-
Tskhali Basin, in the vicinity of Lentekhi and in the sources of the River
Kheledula. Vinogradov (1870) wrote that beavers lived in the bottomland
of Malyi Zelenchuk on the Trans-Kuban Plain.
Bogdanov (1873) reported that beavers were found in the rivers of the
Trans-Kuban steppe, and that the last beaver was killed in 1864 on the
Laba River, not far from its estuary. Dinnik (1884b) recorded the beaver's
presence near the source of the Laba. Keppen (1902) published data on
two beavers killed in 1947 [sic] on the Araks and suggested some interesting
291
possibilities of beaver habitation in the upper reaches of the Araks. Linstow
(1908) published prolific but unsubstantiated data on beavers.
Satunin (1920) used Keppen's and Linstow's surveys in his history of
the Caucasian beaver, observing that if the beaver lived in the Supsa Basin
of Colchis in 1909, it probably survives in some uninvestigated corners of
eastern Transcaucasia and the northwestern Caucasus.
Ognev's survey (1947) on the Caucasian beaver with an appended map of
its former ranges (! ?) added new perplexities for future investigators.
When studying the swamps and rivulets of the Colchis in 1931 and 1944
in connection with nutria breeding, the author did not find gnawed tree
stumps or remains of beaver dams, but was nevertheless convinced that
beavers could have lived recently in the Rion valley from the source of the
Kvirila to the Black Sea coast.
There are rivers (they now have bare banks) near the Tskhaltubo resort
and in many other places in Colchis which would have been perfectly suitable
for beaver.
It is difficult to establish exactly when the beaver disappeared from the
Caucasus, but it was probably in the middle of the 19th century.
South of the Caucasus the beaver was known in the past century in
Mesopotamia and was widespread in Iran. It inhabited the rivers of the
Kyzyl-Irmak system in Turkey where an annual yield of 2,000 pelts (!) was
recorded. The beaver might have lived in the swampy area between Kayseri
and Inesu, south of the middle course of Kyzyl-Irmak in the last half of
the 19th century (Danford and Alston, 1877, 1880).
All this data, published first by Keppen (1902) and then by Linstow (1908),
looks very solid but lacks any scientific confirmation, e.g., figures,
photos and collections. Radde (1866) provides another example of
unscientific speculation: he ''placed the Безауег" on the right tributaries of
the Rion in the highlands of Svanetia in small mountain fissures with
turbulent streams avoided even by the otter.
296 It is difficult to imagine that the beaver could have lived in the central
parts of Asia Minor and Iran considering the absence of water and forests.
It is known that beavers feed on wood of poplar, aspen, willow andbirch, and
it is doubtful that dewberry, sea buckthorn, oleaster and smoothleaf elm,
the main contemporary shrubs and trees growing along the riverbeds in
Iran and Turkey, would have been sufficient food for the beaver. However,
the very rapid changes in landscape of the last centuries are a further
consideration. The recent discovery of beaver remains in the strata of the
ancient Scythian site of Neapolis in the Crimea showed that from the middle
of the first millennium B.C. beavers lived on the Salgir River on a part
of the peninsula that was actually dry and completely woodless (Tsalkin,
1947). With reference to old reports of the beaver on the steppes of the
southern Ukraine, Pidoplichko (1951) reasonably explained that, given the
presence of fluvial forests, the beaver could have lived, and could still
live, in the southern steppe zone, or even in the semidesert zone, on rivers
crossing these zones. His reasoning is useful in validating the earlier
distribution of beaver in Ciscaucasia. On the other hand, it is likely that
the unstable regime of mountain rivers and the dry continental climate
were limiting factors in the uplands of southwest Asia.
The beaver cannot live on a rapid, pebble-bedded mountain river for lack
of shelter and food, even though a forest may grow along its banks. A river
with a steep gradient provides no place for the beaver to burrow, and every
flood will inevitably demolish the beaver dams.
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Nevertheless, the beaver survived intensive orogenic disturbances in
such vast intermontane valleys as the Kura and the Rion, and on submontane
plains. More frequently, beavers resettled in the area after peneplanation
had occurred and subsided.
The history of the beaver on the Caucasus presents a particularly
interesting phenomenon: the morphological-physiological development in
an animal native to warm (Mediterranean) climates of characteristics
adaptive to life under ice.
These adaptations to life in freezing water bodies have often been
emphasized by investigators — Linstow (1908), Fedyushin (1935),
Vereshchagin (1939c) and others — who place their early development in
the Lower Pliocene with the initial cooling of the climate. Swimming under
ice and working incisors under water probably involved a considerably
accelerated process of adaptation in the first half of the Pleistocene.
Postglacial beaver settlement progressed at a startling rate in territories
free from inland ice. In view of the difficulty of migration across
watersheds, the abundance of this animal in historic time in Scandinavia,
on the Kola Peninsula, andonthe Karelian Isthmus might be taken by
opponents of the theory of a major glaciation on the European plain as
substantiation for their views.
The first stage of beaver population decline resulted from excessive
trading in pelts; the second stage and ultimate disappearance of the species
from the Caucasus occurred because of anthropogenic alterations of the
landscape. Environmental conditions changed with the cutting of fluvial
forests and the acceleration of river currents, erosion and flooding as land
was cleared and cattle grazed in the basins. These processes were more
rapid along Ciscaucasian rivers than along those of Transcaucasia. The
beaver survived longest in deep marshes and water courses of the Colchis
lowland.
In the 1940's some beavers were brought in pairs from Voronezh into
the Karayazskii nutria sovkhoz where they reproduced in cages. They could
be allowed to live out of captivity if they were placed in a suitable environ-
ment, e.g., riverbanks grown with poplar and willow trees.
We conclude the following from our survey:
1. The European beaver inhabited the Caucasian Isthmus in the Pliocene
and in the Quaternary.
2. By the beginning of the 18th century, the distribution of this animal
had become vestigial, surviving only in marshes and rivers of western
Transcaucasia, in Colchis, and possibly on the piedmont plains of Cis-
caucasia along tributaries of the Terek and the Sunzha. The beaver is not
known in mountain regions of the Caucasus in historic time.
3. The final extinction of beaver on the Caucasus occurred in the middle
of the 19th or the beginning of the 20th century in Colchis.
4. If water and forest regimes and hunting can be regulated in the
U.S.S.R., it will become possible to breed beavers in small numbers on
some of the left tributaries of the Kuban and on the rivulets of the Khachmas
lowland and of Colchis. Especially suitable for breeding grounds are those
spring-type rivulets ("кага-зи") with a constant flow of water and plantings
of poplar and smoothleaf elm on their banks. Beaver breeding, however,
will not produce any marked effect on an accelerated agricultural economy.
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Family MURIDAE
Remains of 15 species of this family are known from Quaternary deposits
on the Caucasian Isthmus; today the Muridae of the Caucasus comprise
29 species. The emergence of the genus Rattus is of particular interest.
Black rat —Rattus rattus Г. В. cf. rattus remains were known
long ago from Quaternary strata of numerous Western European caves from
the British Isles to the Carpathians (Woldrich, 1882; Brandt and Woldrich,
1887). They were usually found with bones of animals of the Upper
Pleistocene — cave bear, woolly rhinoceros, and others — in Upper
Paleolithic strata, such as Aurignacian strata of the Evelinis cave of
Somerset (England). The majority of black rat bones were found along
Italian coasts and generally around the Mediterranean (Wolf, 1939).
In Lebanese, Syrian and' Palestinian caves black rat bones were found
in the Mousterian strata in which were also found remains of Rhinoceros
mercki and the hippopotamus (!) (Bate, 1937). Black rat remains are, as
a rule, deposited contemporaneously with their enclosing strata, i.e.,
"in situ, '' but there are probably many cases of contemporary rat remains
being thrown down into the bottom of excavations or of the rats themselves
entering the deeper strata by fissures.
Nevertheless, the explanations are insufficient to accept the theory of
postglacial, prehistoric penetration of the black rat by ''importation'' into
Europe — а theory perpetuated in scientific articles. The black rat is
probably a Pliocene relict of the Mediterranean. It survived the Pleistocene
on the warm coasts of southern Europe and north Africa, and perhaps even
on the Caucasus (Vereshchagin, 1947d, 1949c).
In the U.S.S.R. black rat bones were found in Greek and Roman strata
of Olvia, in the southern Ukraine (Pidoplichko, 1938c).
On the Caucasus single bones of
this kind were found in Neolithic strata
of the Akhshtyrskaya cave and in
graves with jar burials of the first
millennium B.C. on the Kura near
Mingechaur (Figure 122).
Actually the black rat is widespread
on the Caucasus, living both ''wild'' and
as commensals with humans. It lives
within the harbor constructions along
FIGURE 122, Black rat pelvis from strata of the the Azov, Black and Caspian sea
first millennium В.С, in Mingechaur coasts. The black rat is not found in
the mountains or plains of Ciscaucasia
and probably has never lived there.
Black rat bones were not observed among the thousands of bones
recovered from eagle owl pellets which were collected by Kistyakovskii
(1935) and the author from the Stavropol and Pyatigor'e areas.
Black rat is widespread along the Black Sea coast from Batumi to
Novorossiisk, living in gardens and forests, and infrequently on the banks
of water bodies where it has been largely replaced by the Norway rat.
Flerov (1927) observed it north of Batumi forest. Shidlovskii (1947, 1948,
1950) recorded the rat in many parts of Abkhazia, Mingrelia and Adzharia.
He claimed that the black rat can be found in Abkhazia from sea level to
600 m and in Adzharia to an altitude of 1,500 m. The author found both
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the black rat and the Norway rat on Bebesyri Lake in the fall of 1946, but
it has hever been observed in the bogs around the Paleostom Lake
(Vereshchagin, 1941b). According to Stepanov (1931), the black rat of
Batumi is less numerous than the Norway rat. In eastern Transcaucasia
it lives ''wild'' in the forests of northern Armenia where it is also found in
populated areas. In 1937 it was frequently observed in the Shamlug mine
and in the mills of the villages of Kulali, Achadzhur and Marts (Shidlovskii,
1942). The animal is rare in South Ossetia and is found only in inhabited
places deepinravines. In Kakhetia andthe Alazan-Agrichai valley it lives in
gardens and buildings, quite often in garrets (Argiropulo, 1940a). It is
common in summer in blackberry thickets bordering fields near Kutkashen
and Vartashen. In the Kura lowland it lives in oases, especially in black-
berry thickets along irrigation ditches.
In Baku the black rat finds refuge in structures along the littoral strip
and usually is not found farther than 100 m fromthe bay's edge. Sixty
percent of the yield of a large-scale capture was from ships in Baku port
and only 9% from port buildings. The remainder of the yield was made
up of the Norway rat (Vereshchagin, 1942a).
This rat is common in the Lenkoran lowland from the village of Masally
to Astara, and further along the southern coast of the Caspian sea in Gilan
and Mazanderan. In these areas it lives in orchards and can be found in
all harbors, especially those near fisheries. Rodents from Asterabad were
known to zoologists even at the beginning of the last century (Ménétries,
1832; Radde, 1886).
According to records from large-scale catches in Lenkoran and adjacent
villages (late 1930's-early 1940's), the black rat comprised only 0.51%
of all synanthropic rodents caught (Vereshchagin, 1949d). Even here the
rodent does not inhabit mountains but confines itself to lower altitudes.
The ability of the black rat to exist in the wild the year round in Colchis
and Asterabad and to construct burrows under tree stumps, fences and
shelters in hollows substantiates its Pliocene origin in the Mediterranean
area in general, and in Transcaucasia in particular.
The black rat adapted readily to the new types of shelter provided by
human culture: first mud huts, and later brick, stone and wood buildings
and boats and ships, in which heat and food were available. Adaptation to
life within harbor constructions and on ships is, in our opinion, associated
with the rat's original habitat on the warm Mediterranean coasts, where
feeding along the shore was possible. Later, penetrating inland along rivers
and confining itself to inhabited places, it gradually occupied the lower
Transcaucasian forests.
Norway rat —Rattus norvegicus Berk. There are no proved
remains of this species from the Pleistocene strata of southwest Asia, the
Caucasus and Western Europe. Remains of Norway rat found in the caves
of Czechoslovakia, Malta, Gibraltar and Ireland were bedded together with
the remains of postglacial animals and the type of preservation does not
indicate an early origin (Wolf, 1939). The bones of Norway rat are known
on the Caucasus from later Middle Ages strata (15th-16th century, A.D.)
of the excavation of the courtyard of the palace of the Shirvan Shahs in Baku
(Vereshchagin, 1949c), and from eagle owl pellets, no more than several
hundred years old, found in small caves in the Stavropol and Pyatigor'e
areas and in North Ossetia and Imeretia.
295
The first reference to mice as being harmful to crops and trees was
made by Claudius Aelianus in the 3rd century A.D. (Latyshev, 1904).
Arabs probably knew of the Norway rat in the early Middle Ages (Zhitkov,
1944).
Nowadays the Norway rat is reported on the Caucasus from all inhabited
places of the country which are connected by railways or roads, from
coastal villages, fisheries and ports along the shores of the Caspian and
Black seas, and from the rivers (Kuban, Sulak, Terek, Kura and Rion),
especially from their navigable sections (Vershchagin, 1947d) (Map 52).
On the Caucasus the Norway rat is frequently found living off the land,
especially in the humid foothills of Kabarda, North Ossetia and the Grozny
Region. Here the Norway rat is often a real pest of gardens and field
storehouses (''sapetki'') (Rossikov, 1887; Bogdanov, 1936). In Kabarda and
North Ossetia the Norway rat lives in mountain villages (''auls''),
particularly in the vicinity of mills, almost to the upper areas of the
Greater Caucasus ravines (Turov, 1926c).
There are wild populations of this rat in the Colchis swamps (Figure 123),
in Asterabad and on ancient lakes of the Kura left bank in Azerbaidzhan
(Vereshchagin, 1914b, 1942b, 1947d, 1949d; L'vov, 1949).
The Norway rat also lives in rock placers on the Caspian desert coast,
e.g., оп the Apsheron Peninsula, where it feeds on flotsam. In the Rion
and Kura lowland swamps, the animal shelters in open, raised nests in
reed thickets or in closed shelters constructed beneath alder roots.
In the wild, the Norway rat feeds primarily on vegetable food but its
consumption of animal food is also significant.
300
FIGURE 123. Habitat of Norway rat in Colchis
Photograph by author, 1939
296
During periods of prolonged snow cover and hard-frozen open waters,
some of the Norway rat population of reed and alder swamps in Trans-
caucasia die in their own habitats, and some survive in the inhabited areas
along the shores (Vereshchagin and Dyunin, 1949).
This species, closely associated as it is with water biotopes, probably
could not tolerate glaciation either in Ciscaucasia or Transcaucasia.
The wider contemporary distribution of Norway rat throughout the world,
as compared with black rat, is consonant with its Pliocene origin, and
with the Mediterranean Holocene origin of the black rat.
The Norway rat possesses a greater adaptability of physiological functions
than the black rat (Ruttenburg, 1950) which has facilitated its settlement
along with man's from the tropics to the Arctic. However, it is less
resistant to winter conditions in the wild in Transcaucasia because of its
hydrophilic nature.
In developing Zhitkov's (1944) opinions and our own main premise of
the survival of the genus Rattus in the Mediterranean from the Pliocene
(Vereshchagin, 1949c), it was necessary to consider the Norway rat as a
postglacial newcomer into the Transcaucasian Swamp fauna and to the
forests of the Ciscaucasian lower mountain zone. This raised questions as
to the routes and dates of its penetration into the Caucasus.
The Norway rat certainly could not have entered Europe from central
Asian deserts. The more recently popularized notion that the species
entered the Caucasus by way of caravan routes across the dry Iranian
desert from India was properly discredited by Kashchenko in 1912. The
301 route of penetration obviously was along the sea coasts. In view of the
historical fact that Egyptian and Phoenician ships crossed the Red and
Mediterranean seas as early as the second millennium B.C., later sailing
as far as southern Asia(Avdiev, 1953), it seems clear that the Norway rat
was transported into the Mediterranean before the common era, probably
first to the coast of the Black Sea in the Caucasus.
The penetration of the Caspian coast occurred later, after settlement
of the Black Sea area, either by water — on the Manych — or by land —
very possibly along the humid, sloped plains of Ciscaucasia.
Before urban concentrations of many-storied buildings and slaughter
houses, and the development of canal, railroad and automobile trans-
portation, the Norway rat apparently lived wild in reed- grown swamps and
river deltas, as canbe seen from observation of its habitats inthe estuaries and
lower reaches of the Danube and the Dniester (Aizenshtadt, 1950b), in the
swamps of the Rion, and in the Volga delta.
Pallas [1741-1811] observed rats in the Volga delta; at that time, the
Norway rat probably lived there, as did the water vole.
The new developmental stage of the Norway rat range in the Caucasus is
associated with the construction of paved roads and railroads, and rural and
urban centers in the 19th and 20th centuries. Slowly it extended into the
populated areas of the Kura lowland along waterways (the Kura and Araks
rivers and their tributaries). The rats first penetrated the Alazan-Avtaran
valley by the left tributaries of the Kura: the Turyan-Chai, Geok-Chai and
Alindzha-Chai. Later the population growth was accelerated by road
construction — less so on the dry Armenian plateaus than elsewhere. In
Yerevan and Dzhulfa, for instance, the Norway rat appeared inthe 1930's after
the construction of the railroad connecting these towns with Tiflis and Baku.
297
302
In the Ciscaucasian Plain the settlement of the Norway rat accelerated
in the 1920's with the development of road transport. Gubarev (1941)
recorded the appearance of the Norway rat in inhabited areas of the Salsk
steppes.
The Norway rat was known to Satunin (1901) in Stavropol from bones
contained in eagle owl pellets from Mount Strizhament, which established
that the species inhabited the forested valleys of this region as early as
the last century.
Kuzyakin's survey (1951) on the Norway rat's origin in northern Eurasia
and its relationship to man added nothing to an understanding of the history
of the animal in the U.S.S.R.
Striped field mouse — Apodemus agrarius Pall. Remains of the
striped field mouse are known from Pleistocene strata of Czechoslovakian
caves (Wolf, 1939). They are as yet unknown on the Caucasus, although
they have been exposed in Upper Quaternary deposits of the lower Don.
The contemporary range of the striped field mouse occupies the broadleaf
forest zone and the southern Eurasian taiga with discontinuities in Trans-
baikal and in the upper Amur Basin (Bobrinskii, Kuznetsov and Kuzyakin,
1944). The striped field mouse inhabits only the river valleys of the Russian
Plain steppe zone. It is now absent from the Crimea although it has been
found in Pleistocene strata. The distribution in Ciscaucasia, now isolated
from the north, is confined to the humid piedmont and to the riverbeds of
the steppe and semidesert (Sviridenko, 1944). Striped field mouse has
been reported from the Black Sea coast south as far as Sukhumi (Shidlovskii,
1947, 1950).
The species inhabits the Stavropol Plateau and the tugai forests of the
Kuma, Terek and Sulak rivers. An isolated range is situated to the southeast
in the forests of the Khachmas lowlands from the lower reaches of the Samur
to Khachmas (Beme, 1928; Vereshchagin, 1944).
Its vertical distribution is insignificant: the upper parts of inhabited
mesophytic meadows and fields in the northwest Caucasus between altitudes
of 950-1,000 m; inthe Nalchik area, to an altitude of 700 m; and near
Ordzhonikidze and in Alagir, between 800 and 850 m.
Sviridenko (1944) observed that striped field mouse distribution is related
to the amount of precipitation. The animal becomes eurytopic only if the
annual precipitation is above 500 mm. In more arid regions the rodent
inhabits only humid biotopes like the lowland forests of the Khachmas
depression near the Caspian coast. Here the striped field mouse lives in
the humid bottoms of afforested valleys and on the banks of spring-fed brooks
under cover of alder and blackberry growth interspersed with liana. In this
environment the mouse is not harmful to cereal crops as it is in other places.
On the more arid Kusary sloping plain and on the remainder of the Caspian
coast, this animal is not encountered.
The distribution of striped field mouse in Ciscaucasia is mostly confined
to regions with a dense network of small torrents and springs, such as the
developmental zone of piedmont trains. Shrub areas along small rivers
and small meadows along turbid brooks are characteristic biotopes.
Consequently, the Caucasian foothills contain several isolated ranges:
the Kuban (including the Black Sea coast), the Terek-Sunzha and the Samur
(Map 54). The breaches in the Ciscaucasian range areas are created by
dry, steppelike divides.
298
Farlier — in the Pleistocene — the striped field mouse barely penetrated
farther south than the lower forests of Khachmas, Since its remains were
not found in the Pleistocene asphalts of the Apsheron Peninsula. This is
further substantiated by its absence from the Alazan-Agrichai valley where
conditions are suitable for its habitation.
In humid western Transcaucasia the striped field mouse is distributed
along terraces of the Black Sea coast southward to Sukhumi. Here it is
frequently observed in glades, clearings, shrubs and orchards.
Shidlovskii's data (1947) (Table 70) are characteristic for striped field
mouse distribution inthe biotopes on the Black Sea coast.
TABLE 70, Distribution of the striped field mouse according to biotopes in Abkhazia
Vegetable
Orchards
plantations
Com fields
reservoirs
Continuous
Total
Number of (absolute
animals
1 27 36 12 10 31 135 2
12.2
number
in %) ..
caught
303 These data show that anthropogenic alterations of the topography in the
Black Sea littoral zone, especially deforestation, favor the growth of the
distribution and population of the rodent. It is possible that the considerable
extension of the striped field mouse range south along the Black Sea coast
in historic time was promoted by man-made changes in the landscape.
However, farther south in Colchis, important to note, there are no
occurrences of striped field mouse, although the biotopes are much like
those of Abkhazia, a fact probably explained by the excessive humidity.
The peculiar underdevelopment of the field mouse ranges on the Caucasus
suggests that the penetration of this species from the north occurred rather
late in the Pleistocene. With the postglacial advance, the Caucasian
populations of the striped field mouse were apparently cut back by the
development of a large belt of dry steppes.
The absence of the animal from the Crimea in the Recent and in the
Pleistocene also confirms a late development of the range in the south.
In the Recent epoch, the striped field mouse's distribution on the Caucasus
increased in the southwest with forest clearing in places of excessive
moisture, and in the dry regions to the north and south with the development
of irrigation.
Asia Minor hamster — Mesocricetus auratus Water. (5. lato). The
question of the origin of the hamster of genus Mesocricetus on the
Caucasus is rather complicated, for the earliest hamster remains —
genus Paleocricetus — found on the Caucasus date from the Middle
Miocene. In the Lower Pliocene of Ciscaucasia there lived a hamster
almost the size of the golden hamster of the eastern Mediterranean area.
Thus, the Caucasus could have been as much of a focus of Asia Minor
(golden) hamster development as was southwest Asia.
299
304
Fragments of hamster remains of the genus Mesocricetus are known
from the Middle and Upper Paleolithic of Palestine (Athlit cave; Picard,
1937) and, within the limits of the Caucasus, from the caves of the Rion
Basin and Kvirila River (Gvardzhilas, Mgvimevi, Kudaro) and from
Pleistocene strata of the Apsheron Peninsula.
It is important to emphasize that, until now, the only hamster remains
found in Quaternary deposits of the Russian Plain have been those of the
genus Cricetus.
Postglacial cave deposits of Asia Minor hamster remains are quite common
on the Armenian Highland, in central Transcaucasia and in central
Ciscaucasia.
In recognition of its considerable geographic variability, the Caucasian
representatives of the genus Mesocricetus are classified by
taxonomists under two or three separate species. Accepting southern
southwest Asian origin of the extant Caucasian golden hamster, we can
further assume that, in chronological order, Mesocricetus,
M. auratus raddei Nehr., and M. auratus nigriculus Nehr.
comprise a genetic series of successive subspecies.
The contemporary range of this hamster includes Syria, Asia Minor,
the Armenian Highland, Talysh, the northwestern Iranian Plateau, eastern
Transcaucasia, central Dagestan and the Ciscaucasian Plain. Surveys of
the distribution areas were given by Argiropulo (1935, 1939a), Neuhduser
(1936), Shidlovskii (1940c) and Ellermann (1948). The vertical distribution
of Mesocricetus lies within a range of 0-2,400 m. The altitudinal
optimum of the Transcaucasian subspecies M. auratus koenigi Sat.
is, according to Shidlovskii (1940c), in the range of 1,200-2,000 m (Map 58).
On the Armenian Highland within the limits of the U.S.S.R., this
hamster is widespread in relatively warm and arid upper river valleys.
It inhabits the festuca-grown upland steppe of the Pambak River valley,
the borders of Lake Sevan, and the Saraibulakh and Zangezur ridges. On
the cold upland of Dzhavakhetia it can be observed on the steppe meadows
near Tabistskhuri Lake.
In the Karabakh uplands the hamster is known from the Gevorkevan
vicinity.
In Talysh it has been encountered in an area stretching from the upper
reaches of the Vilyash-Chai south to Kel'vyaz, in the upland zone of
artemisia—grass-steppe at an altitude of 1,600 to 2,000 m. It was not found,
however, in groupings of small goat's thorn and acanthus as in the
Armenian Highland but, rather, concentrated in more humid places in the
valley bottoms, especially on the margins of sown fields.
The animal's distribution on the ranges of the northern Iranian Plateau
is probably discontinuous.
In eastern Transcaucasia the hamster inhabits the Gori depression,
occupying regions of earlier steppes or forest clearings. In the Pleistocene
and Holocene, it was distributed in Imeretia and in the upper reaches of
the Rion. Eastwards, onthe Iora and Kartalinia plateaus, the animal
inhabits areas grown with the racemose andropogon at altitudes of
550-600 т. It is not found east of the Alazan to Shemakha, but it is possible
that relict colonies will be found northeast of Shemakha, for example, in
the Kilyazi-Chai and Gil'gin-Chai valleys, and west of Derbent.
300
305
The distribution of the Dagestan subspecies, М. auratus raddei
Nehr., is limited in inner Dagestan by the valleys of the Argun, the
upper Sulak and the Samur. The animal has been traced from the village
of Shatili to Evdokimovskoe in the Argun valley. According to Rossikov
(1887) it is known from the Andi KoisuRiver. Accordingtothe data from the
Orlov expedition, Beme's records (1925), and Dyukov (1927), the hamster
is encountered on the whole of the Khunzhakh uplands on the fields
surrounding Baitl, Chondotl and other villages.
Here the hamster inhabits the mountain xerophytic steppe andis attracted
to fields of grain. Its vertical placement is from 1,600 to 2,300 m (Geptner
and Formozov, 1941).
In the Middle Pleistocene these ranges were probably linked with the
piedmont areas of the Apsheron.
Perhaps this link was broken in the Upper Pleistocene when Dagestan
was divided by water erosion and the formation of a forest zone on its
southeastern border.
The present-day range has enlarged somewhat under the influence of
agricultural activity.
The distribution of the Ciscaucasian form M. auratus nigriculus
Nehr. is very extensive, although always ''squeezed'' close to the mountains
(see Map 58).
Its habitat in northern Ciscaucasia is in the zone of artemisia—festuca-
steppes, while on the Stavropol Plateau and in the Kabarda foothills it is
found on steppe-meadows and plots planted to corn. The animal is also
encountered in the motley-grass of the Terek valley (Ognev, 1924).
In many places this hamster has actively penetrated the mesophytic
foothills of Ciscaucasia where the forests have been cut, for instance on
the Trans-Kuban, Kabarda and Terek-Sunzha sloping plains. In general, this
Ciscaucasian form is more eurytopic than its Transcaucasian parent,
although, in the main, it occupies dry places.
Its vertical distribution in Ciscaucasia normally extends from sea level
to 1,000 m. In central Ciscaucasia, it penetrates the mountains as far as
Kislovodsk and Nal'chik, but is not found in dry, longitudinal valleys of the
northern slope. It is spread along the valleys of the left tributaries of the
Terek. M. auratus nigriculus is distributed only as far as the
Chernye Gory canyons.
In the last few decades the settlement of hamsters in the Rostov Region
has been reported and in the 1940's they were found northwest of Manych.
Judging from the Asia Minor hamster's contemporary range and гот 115
level of trophic succession, it seems possible that the ancestral species
penetrated the Caucasus from the south and southwest during the Tertiary,
most probably during a dry phase of the Upper Pliocene.
The path by which the hamster extended its range from the Lesser
Caucasus to eastern Ciscaucasia probably passed the foothill steppe zone
along the base of the southeastern slope of the Greater Caucasus and farther
through the present uplands of Dagestan which at that time were lowlands.
Later this route was impeded by loam and pebble sediment on the swamped
plains of the piedmont, particularly in the region of presently eroded
sections of Adzhinour and of the Kusary sloping plain. The distribution area
of the hamster was probably the widest in the Middle Pleistocene during
the wide steppe formation on mountain slopes. The subsequent mesophytic
trend in landscape and the shifts in altitudinal zones during glaciation
301
306
created а break and caused the decrease of the hamster range. Another
breach in the range occurred with the onset of the postglacial xerothermic
period which produced a discontinuous upland mountain relief in southwest
Asia. N
On the other hand, in the humid mountain regions of Imeretia and South
Ossetia the Lower Holocene desiccation of ridges and slopes contributed
somewhat to the spreading of the hamster range; there it was only displaced
later during another humidification of the climate.
There can be another interpretation of the history of the hamster range
development on the Greater Caucasus and in Ciscaucasia, if, while accepting
two distinct species for the Dagestan and Ciscaucasian forms, we consider
the possibility of their having a common ancestor in a local Tertiary form.
In the Recent epoch, ranges preserved in southern Transcaucasia are
relatively stable and are diminished only during cattle grazing, large
scale plowing of artificially watered land, or purposeful killing of the
animal as a pest through the use of chemical exterminators.
Asia Minor gerbil — Meriones blackleri Thos. Remains of gerbils
of the genus Meriones were recorded from Acheulean and Mousterian strata
of Palestinian caves (Picard, 1937; Wolf, 1939). In the Caucasus, aside
from findings of Gerbillus sp. in Pliocene strata of the Armenian
Highland (Bogachev, 1938c), only the remains of the present-day species
are known from the superficial layers in caves and shelters of the Araks
valley.
The present-day rangeof Meriones blackleri Thos. includes
Asia Minor, Syria, western Iran and a part of Transcaucasia (Ellermann,
1948; Bobrinskii, Kuznetsov, Kuzyakin, 1944; Pogosyan, 1949).
In Transcaucasia Meriones inhabits the middle Araks valley, certain
places in the Araks gorge in the Zangezur ridge, the eastern Karabakh
foothills, the Mil'skaya steppe, the southern Mugan steppe and the Kirovabad
sloping plain. In the wide valley of Akera-Chai, it is found almost as far
as Lachin, that is, about 1,500 m above sea level. The population of
Meriones blackleri Thos. inthe plains of the middle Araks Valley
is greater than that of other species; on the stony slopes, however, it is
superseded by Meriones persicus Blanf. The animalis not encountered
on the left bank of the Kura, although conditions are the same there
as on the right bank. On the Kirovabad sloping plain its distribution can
be traced to the lower reaches of the Khram. According to data from large-
scale catches, it has a greater population than Meriones erythrourus
Gray in the foothills of the Lesser Caucasus.
In the Mil'skaya steppe М. blackleri can be seen with М. егу-
throurus almost everywhere, but in the Karabakh foothills it is
encountered only in warm areas with artemisia groupings at altitudes of
350-400 m above sea level. In the northern part of the Mugan steppe, this
animal has been largely dislodged by cultivation and irrigation and is seen
mainly in artemisia semidesert along the border from Karadonly to
Belyasuvar. Оп the northeastern slopes of Talysh, it has been traced to
the Eshakchi outpost at altitudes of about 600-700 m (Map 63).
The animal mainly inhabits alluvial and talus deposits of loessial
loam in large fluvial valleys grown with artemisia and ephemeral grasses.
In view of M. blackleri's specific distribution in eastern Trans-
caucasiaand, more particularly, its absence from the Kura-lora interfluvial
302
307
area and the Shirvan steppe, we can assume that it settled in the north only
relatively recently, i.e., in the’ Pleistocene or perhaps even in the
Holocene. In southern Transcaucasia in the middle Araks valley,
M. blackleri may be considered a local species fromthe Upper Tertiary.
Persian gerbil— Meriones persicus Blanf. Fossil remains of
this species from Pleistocene strata have not been validated, but they are
known from Holocene cave deposits on the southern slopes of the Armenian
Highland and from the middle Araks valley.
The main part of the M. persicus range (Map 62) includes Turkey,
Iran, Afghanistan and southern Turkmenia (Neuhauser, 1936; Geptner,
1940; Ellerman and Morrison-Scott, 1951).
On the Caucasus it is eurychoric over the southern slopes of the
Armenian Highland and in the highlands of Talysh. Its population in the
Alindzha-Chai Basin, according to large-scale catches, comprised 62.5%
of the total Meriones population. It is not encountered on the left bank
of the Araks gorge north of Mindzhevan. Inthe gorges of the southern
slopes of the Zangezur ridge, the range reaches an altitude of 1,800 m in
astragali_tragacanth formations and stretches as far as Zangelan and
Shakhbuz.
On the Saraibulakh ridge, according to Dal' (1904b), Meriones
comprised 17% of the rodent constituent of the food of the eagle owl. It is
observed in the Zanga valley to an altitude of 1,230 m (Dal', 1948b) and
is common on the slopes of the outlying northern ridges of the middle Araks
valley among mountain halophytes of the semidesert (Flerov and Gureev,
1934; Argiropulo, 1939b; author's data, 1947).
In the highland steppe of Talysh the species is common in the upper
reaches of the Vilyash-Chai and Zuvanda at altitudes of 1,500-1,800 m.
Our data collected in the vicinity of the Dyman outpost in 1945 show that
its remains constituted 26 % of the remains of all rodents found in owl
pellets.
In the dry mountain areas ofnorthernIran and Asia Minor M. persicus
is evidently the most numerous and eurychoric. Its usual biotopes are the
stony slopes of valleys which are dotted by cushions of thorny astragali,
acantholimon and artemisia, and in spring by ephemeral grasses and
poppies. In certain places the animal emerges on divides beneath the
mountain-grass and steppe zone.
The facility of М. persicus in digging its burrows under stones or
shrub roots, its skill in climbing stones, its hairless feet, long tail with
panniculus and its long vibrissae —all these features, according to
Argiropulo (1939a), are early adaptations to a mountaindesert habitat.
This animal is mainly granivorous: during early summer, it feeds
mostly on grain of small grasses; in later summer, on grain of mountain
saltwort. It is an early indicator species of the southwest Asia mountain
desert mammal complex.
M. persicus thrives in agricultural regions. In grain fields and along
bordering fences, its population is always increased somewhat. Cattle
driving and overgrazing contribute to the spreading of thorny astragali, thus
extending the range.
Red-tailed Libyan gerbil— Meriones erythrourus Gray ($. lato).
Fossils of this animal are known from Middle Pleistocene strata of the
Apsheron Peninsula.
303
308
At present, this polymorphous species (for which as many аз 23
subspecies have beenrecorded) is identified with M. libycus Licht. Its
vast range includes north Africa, southwest Asia, eastern Transcaucasia
and central Asia to Semireche [Dzhety-Su]. Its northern limit on the
Caucasus passes the latitude of Shemakha (Ellermann and Morrison-Scott,
1951; Bobrinskii, Kuznetsov, Kuzyakin, 1944).
In Transcaucasia it inhabits all of the Kura-Araks lowland and foothills —
from Tiflis on the west to the Caspian coast on the east (Map 61) — and
is especially numerous on the Apsheron Peninsula. М. erythrourus has
not been observed north to Kilyazi or south to the Kyzyl-Agach Gulf on the
Caspian coast.
The species is encountered throughout the central lowland. A dense
-population is noted among artemisia, saltwort and caper formations in the
flat semidesert of the Mil'skaya, Shirvan and Mugan plains. The animal
settles more frequently on alluvial ranges and on the margins of minor
solonchak depressions overgrown with arboreal saltwort shrubs and
Austrian and Sovichwormwood. The maximum population is observed,
however, on the sandbank ridges of the Kura, Alazan, Гога and Araks,
especially in the tamarisk-thicket zone. Similar high-density populations
are encountered on the dunes of the Caspian coast where the sand hillocks
are held by tamarisk and ephedra. In such places there are perhaps
40 specimens per hectare. Biotopes created by man, especially vineyards,
abandoned irrigation systems, shepherds' camps in the steppes and fallows
of several years standing, are of great importance tothis animal's settlement.
The animal is crepuscular and nocturnal in summer, crepuscular and
diurnalinspring and fall, anddiurnalinwinter. Meriones erythrourus
is mainly granivorous: in spring it feeds on shoots of ephemeral annual
plants; in summer, on shoots and grains of goatgrass, ragweed, brome
and insects, in particular on non-gregarious locusts; in fall, on new shoots
of ephemeral grasses; in winter, on wormwood stalks, seeds of arboreal
saltwort and others.
In its search for grain, the animal climbs high on the saltwort shrubs.
During snowfalls these animals usually lie in burrows untilthe thaw, rarely
grazing on adjacent shrubs. The abundance of animals on sand hillock
ridges in level steppes is explained by the ease of constructing ventilated
burrows and the opportunity of winter fattening on artemisia and saltwort.
The coincidence sometimes seen of dense rodent populations in cattle camps
(''yatagi'') in these areas might persuade an unskilled observer that cattle
raising is favorable to hibernating populations of Meriones (Gladkina,
1952).
The contemporary range of M. erythrourus and its adaptation
to the landscape indicate that the species formed in the southern and
southeastern Mediterranean range. It penetrated the Caucasian Isthmus
from the south and southeast probably as early as the Pliocene but no later
than the Lower Pleistocene, since by the Middle Pleistocene it already
inhabited the Apsheron Peninsula.
The animal survived the transgressions of the Caspian Quaternary on
the Tertiary hillocks on ihe northern and western borders of the Kura Bay.
In the eastern part of the Kura-Araks lowland, this species is one of several
late settlers of the territory after the sea waters retreated from it in the
Holocene.
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Local agriculture has, untilnow, been favorable for the development of
the species. However, the construction of irrigation works and the expansion
of irrigated cotton fields will rapidly drive this animal from vast land areas.
Steppe vole —Microtus socialis Pall. (s. lato). Remains of this
genus have been found in Middle Pleistocene strata of the Apsheron
Peninsula, in Holocene cave strata of the Saraibulakh Range in Armenia
and in many other places on the present-day range, which includes
Cyrenaica, the Balkan Peninsula, the southern Ukraine, the Crimean steppes,
southwest Asia, central Asia, eastern Ciscaucasia and eastern Transcaucasia
(Minin, 1938; Vinogradov and Argiropulo, 1941; Ellermann, 1948).
In Ciscaucasia this vole inhabits the Salsk steppes along the Manych
valley, the regions of the lower Malka and Baksan, the slopes of the Terek
and Sunzha ranges and steppelike piedmonts of northern Dagestan. The
animal penetrated the Terek-Kuma massif of the sandy semidesert along
the network of ducts and channels of the Kuma, Kura and Terek.
On the sloping plain of the Kusary, the steppe vole's range is found in
warm places to an altitude of 600 m, adjacent to that of the common vole.
The animal inhabits only wide plowed fields in lowland forests of the
Khachmas plain.
From the western Caspian coast the range extends into eastern
Transcaucasia at altitudes of 700-800 m on piedmont ridges (Map 72).
The vole penetrated the lowland forests of the Alazan-Agrichai valley
through deforested areas. On the Apsheron Peninsula and in Kabristan,
there are small relict colonies in places with a maximum annual
precipitation of only 280 mm, relieved, however, by localized, supplemen-
tary moistening of the soil and an underlay of impervious layers of
Apsheron limestone. The continuous distribution of the animal in the Kura-
309 Araks lowlands begins in a region where the annual precipitation is less
than 350 mm. In southeastern Mugan the distribution toward the Lenkoran
lowlands is limited by greater soil moisture in winter and 800 mm annual
precipitation (Vereshchagin, 1942c, 1946Ъ). *
In the Kura valley the vole was found only as far as the Mtskheti until
recently, when it penetrated the Mukhran valley as a consequence of
deforestation there. Near the Mtskheti the range is broken by the forested
narrow gorge of the Kura, and resumes again in the dry Gori depression
at altitudes of 600-800 m. Here, M. socialis goriensis Arg. was
isolated as a subspecies, slightly different from the typical species. The
vole inhabits the Kirovabad sloping plain up to the existing lower timberbelt,
i.e., to 600-700 m, although in some gorges it is encountered even higher.
On the deforested northeastern slopes of Karabakh, the steppe vole has
spread upward to 800 m in the vicinity of Madagiz, Mardakert, Martuni
and other points where its range is adjacent to the lower distribution belt
of the pine vole. The vole penetrated high into the mountains through the
valleys of the Akera and Okhchi-Chai via the southern slopes of Karabakh.
A similar penetration can be observed in gorges of the southern slope of
the Armenian Highland.
In the semidesert of the Kura-Araks lowlands the steppe vole forms
temporary settlements in caper formations on chestnut soils, and lesser
settlements in the artemisia—grass groupings. Summer conditions for the
animal in the semidesert zone are rather poor because of lack of moisture
* К.М. Rossikov's report on the mass daytime migration of voles during August in the Mil'skaya Steppe,
noted by Ognev (1950) is, in fact, only a fiction (Vereshchagin, 1946Ъ).
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310
in food and the high temperature of the soil. The vole withstands these
conditions by feeding partially on locusts and mollusks and by constructing
deep burrows (Vereshchagin, 1946b).
The semidesert population of voles declines catastrophically each year
as June approaches because of cessation of reproduction and the sudden
death of mature animals brought about by the desiccation of food and the
high temperatures. Viable isolated colonies of animals survive in some
hollows, in ravines grown with milk thistle (Silybum), in gardens
moistened by irrigation, or under caper brush, etc.
In the Mugan steppe, in the area of Belyasuvar and Astrakhan-Bazar,
the years of peak breeding witness haphazard nocturnal mass shifts of
the steppe vole in the first half of June because of physiologic disturbances
aroused by the desiccation of vegetable food and soil overheating (Mamedov,
1950). *
Because vole behavior shows seasonal peculiarities controlled by
temperature and insolation, in Azerbaidzhan this animal is more vulnerable
to attack by predatory birds and mammals during the winter.
Characteristic ''pulsations'' in the ranges of this species can be traced
over the years near the base of the Apsheron Peninsula. When winters are
warm and humid and summers cool for several consecutive years, there
is an acceleration in reproductive activity and a significant increase in
population. In the semidesert when the fall is humid, large food reserves,
consisting of plant shoots and motley-grass—ephemeral grass formations
are built up. This makes it possible for the animals to move from the
Kabristan ridges through drying rivulet valleys almost to the sea coast,
30-40 km beyond the outer limits of their fixed habitat.
Meanwhile the first hot and dry season again causes mass mortality
among the animals and a new withdrawal northwestward from their
distribution area. This phenomenon is very significant in explaining the
mode and rapidity of migration and the history of the animal range areas
resulting from secular climate and landscape variations.
In Talysh the vole may be found at altitudes of 1,500-1,800 m in the
grassy mountain steppe, while it is absent in the poor fodder grouping of
thorny astragali. A dense vole population developed on the eastern and
northern slopes of Talysh near the upper timberline, which has been
considerably lowered because of grazing andcutting. The upper parts of the
Vilyash-Chai gorge inthe Yardymly region and the area of Vassaru-Chai
near Lerik which is planted to grain are the real foci of mass vole
reproduction. Such foci are the results of human activity. From them,
the vole penetrates downwards to the gorge bottoms of the eastern slopes
of Talysh only whena forest has been thinned or completely cleared.
In the middle Araks valley the vole is found in Aralik and on some of
the tributaries of the Araks, such as Alindzha-Chai, Nakhichevan-Chai
and Arpa-Chai. The vole is not observed on the sun-scorched slopes and
ridges ofthe Zangezur Range inthe zone of stony-mountain—saltwort-semi-
desert and tragacanth astragali because of the poor nutritive value of the
vegetation. In the central Armenian Highland the vole penetrated into the
Araks valley by the left river bank. A steppe vole habitat is located in the
upper part of the Pambak River valley, in the grassy mountain-steppe zone,
* Ognev's report (1950) on the habitat of this vole in the Lenkoran lowlands is based on a confusion in the
location of geographic points: the Kelvyaz outpost is in the high Talysh Mountain and not in the Talysh
lowlands,
306
at ап altitude of 1,200-1,800 m. This site is cut off from the west by the cold
Akhalkalakhi Plateau, and from the north by the afforested spurs of the
Bzovdal'skii ridge and the Pambak River gorge. Contrary to Ognev's
opinion (1950), the range of this species bears no relationship to that of the
Gori subspecies. Such a relationship should be sought to the south in the
Leninakan area and Kars Upland where the dry mountain valleys probably
have populations morphologically similar to M. socialis schidlovskii.
Dal' and Zakharin (1951) indicate that this vole's distribution extends from
the upper reaches of the Pambak through the Araks valley up to the latitude
of Yerevan.
The population of the steppe vole is greatest during the reproducing
season, when the number of burrow openings observed per hectare attains
90,000 on both plowed and unplowed land in the piedmont andropogon-steppe
zones of the Adzhinour and Kartalinia plateaus, the eastern foothills of
Karabakh and the northwestern slopes of Talysh. In this steppe landscape
the contemporary ecological optimum of the species has been reached
(Satunin, 1912a; Vereshchagin, 1942c, 1946b).
Viable even in periods of strong inhibition of population, the colonies of
this rodent are clearly associated with shrubs of Christ's-thorn which
affords them a safe shelter from predatory birds and extreme heat. The
juniper— pistachio forests adjacent to the beard-grass steppe — the so-called
"srid stunted forest'' — is sparsely populated by voles; they are encountered
in thinned forest glades of motley-grass where they construct burrows under
ephedra shrubs.
The characteristics and the periods of steppe vole reproduction are
dependent upon altitude and man-made landscape alterations. In the
semidesert zone of the eastern Transcaucasian lowland, its period of
reproduction is confined mostly to October-November and to March-April;
although if the winter is warm and ephemeral vegetation available, it may
extend from October to May. But in the mountain steppe of Talysh and on the
Armenian Highland, intensive reproduction is observed in June, because
of the later burning-off of the vegetation.
The vole maintains longer sexual activity and vitality on unirrigated
cereal croplands in the piedmont (Figure 124) and on irrigated cereal and
alfalfa croplands in the semidesert zone of eastern Transcaucasia, than
on virgin soil.
Climate is the main factor affecting population variations and area of vole
ranges, especially in eastern Transcaucasia. Nevertheless, epizootic
factors, predatory mammals and birds also figure significantly in reducing
the population of this rodent during peak breeding years. The greatest effect
of bird predation on the vole population can be seen on the semidesert and
the piedmont steppe from November to April, i.e., during the migration
and wintering of the birds (Vereshchagin, 1946b).
Agriculture contributed to the expansion of the range of this species,
opening new territories by the felling of lowland forests, and providing
for an increased population by supplying a succulent and high calorie diet of
cereal crops.
The main morphological features of steppe vole (permeable fur
covering, short extremities poorly adapted to rapid and remote migrations,
flat skull, elongated diastema indicating its enormous burrow-digging
capacity in heavy and medium soils) testify that the species was formed
under topographical xerophytic conditions of the eastern Mediterranean.
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FIGURE 124, М. socialis ina barley field
Photograph by author, 1939
The steppe vole of the Caucasus and eastern Asia never emerges in the
desert proper but is dispersed in its eroded interzonal fringe areas,
indicating an earlier association of presently separated steppes and semi-
desert areas, probably during certain phases of the Pleistocene. The
reasons for this association have been discussed elsewhere by us
(Vereshchagin, 1942c; 1946b) and by Geptner (1945).
Pidoplichko's opinion (1951) on the recent development of the vole range
in the eastern Mediterranean rests on a patently erroneous description of
the distribution area.
The initial evolution of a variety of southern gray voles probably took
place in the eastern Mediterranean on the relatively dry Pliocene uplands
of Asia Minor. One of these forms, the Russian steppe vole, populated the
dry coasts of eastern Mediterranean basins in the Lower Pliocene.
The history of the development of the vole's range on the Caucasus
follows orogenetic movements, fluctuations in the Caspian Sea level and
shifts in landscape zones — both horizontal and vertical — and reveals the
lability of the species in adapting to its environment. For instance, isolation
of the Gori subspecies was caused by the rise of the northern spurs of the
Trialet and the southern spurs of the Kakhetian ridges which partitioned off
the Kura valley near the Mtskheti. M. socialis schidlovskii may
be considered a special species having an earlier isolation. This isolation
was accomplished by the uplift of the Armenian Highland in the Quaternary
which spared some xerophytic communities in separated areas. Perhaps
vast lava flows overlapping the sections of the ancient plateau in different
directions contributed to this isolation (Vereshchagin, 1942c; Shidlovskii,
1945).
The range of the steppe vole on the Isthmus evidently fluctuated repeatedly
during the Pleistocene, conformingto the development and reduction of
landscapes suitable for habitation during different climatic phases. In our
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313
time, forest clearing and slope erosion have facilitated the penetration
of the species into the foothills and mountains, thereby greatly expanding
its range. This is especially conspicuous on the mountain slopes of
Karabakh and Talysh. The vole penetrates the semidesert and desert of
eastern Ciscaucasia when channels are dug and new regions are developed
for grain growing. New extermination methods have no noticeable
reversible effects upon these incursions.
Common vole — Microtus arvalis Pall. The fossil remains of this
animal are known from many Pleistocene cave burials and from alluvial
deposits inEastern Europe, from Ireland to the Carpathians and from Italy
to Sweden. Inthe U.S.S.R. the remains of this vole are common in Upper
Paleolithic strata of the Crimea (Vinogradov, 1937b) and in Middle and
Upper Pleistocene strata of the lower Don and the Urals (Vereshchagin and
Gromov, 1952).
On the Caucasus the bones and skulls of this vole were found in Middle
Pleistocene strata of the Apsheron Peninsula together with those of the
steppe vole (Argiropulo, 1941b; Gromov, 1952). *
In the Recent epoch this vole inhabited the taiga, the forest steppe and
even desert areas from the Atlantic Ocean to the Khingan and the upper
reaches of the Amur. It has a peculiar distribution on the Caucasus
(Map 71).
In Ciscaucasia an almost unbroken distribution range runs from the Don
steppes through Manych along the Azov shore and includes the lower reaches
of the Kuban and the Taman Peninsula.
This vole is widespread in the Stavropol uplands where it lives in
mesophytic meadows and deforested areas.
The northeastern limit of the M. arvalis range in the Ciscaucasian
steppes approaches the longitude of Budennovsk, coinciding with the margin
of semideserts in the lower Kuma area. Eastward it pushes far through
the river valley to the Terek-Kumsa sands. Following the courses of the
Terek and Sulak rivers as far as the Caspian coast, habitats of this vole
occupy ridges and depressions with meadow, tugai and steppe vegetation.
On the Ciscaucasian dry steppe, freshwater basins are a requirement for
the maintenance of vole habitation (Naumov, 1948). On the Greater
Caucasus this vole inhabits only the Trans-Kuban sloping plain in the
foothills at an altitude of 900-950 m, and is not encountered in higher forests.
In the mountain meadows of the Greater Caucasus, it is always replaced
by the pine vole.
The vole is common on the sloping plain of central Ciscaucasia, but is
rarely found, if at all, in the mountains (Ognev, 1950). On the deforested
sloping plain of Terek-Sunzha voles are numerous and apparently
destructive. Turov (1926c) considered the species to be common only in the
forest zone of North Ossetia. According to our observations, the deforested
glades and subalpine meadows in the Fiagdon and Gizel'don gorges of the
Lesistyi and Skalistyi ridges are heavily populated by this animal. It is
common in the Urukh ravine at altitudes of 1,500-1,800 m, and, farther
east, has been traced in the Argun and Sulak basins at altitudes from 800 m
to 2,700 m (Beme, 1933; Geptner and Formozov, 1941).
It has not been observed in the Dagestan foothills from Khasavyurt to
Derbent, or farther southward on the low littoral terrace down to Baku.
It is also absent from the humid regions of the Khachmas lowland which
is inhabited by steppe voles, field and house mice. On the Kusary sloping
* Lower jaws of this vole species were found in the Lower Pleistocene strata of Kudaro I in 1958.
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plain, the lower border of the range is at an altitude of 600-650 т
in the pasture-forest zone, and only on meadow sections of the Samur
valley does it descend to 250-300 m.
If accurate, Beme's mention (1928, p. 143) of animals caught near the
Samur estuary is noteworthy.
At the eastern end of the Greater Caucasus this rodent is encountered
in grain fields and on subalpine meadows in the Konakhkent and Khizy
regions. Its lower range in the Shemakha area extends along an altitude
of approximately 800 m, while farther west on subalpine meadows in the
upper reaches of the Gerdyman-Chai the species is replaced by the
pine vole.
In the deforested areas on the southern slope of the Main Range, the vole
penetrated from mountain meadows through glades overgrown with bracken
by way of the Demir-Aparan-Chai, Bum-Chai and Mazym-Chai ravines
(in the vicinity of Kutkashen, Vartashen and Lagodekhi) up to the base of
steep slopes and into the zone of lowland forests of the Alazan-Avtaran
valley. On the high mountain meadows of the eastern half of the southern
slope ofthe Greater Caucasus, the species is usually concentrated along the
margins of cattle stands heavily covered with manure and overgrown with horse
sorrel andorchard grass. It also populates subalpine meadows on the deforested
ridges of the southern lateral ranges.
The vole is common on meadows of the Kakhetian Range and in intensely
deforested South Ossetia down to the reaches and tributaries of the Rion;
here it is replaced by the pine vole as on the northern slope.
There is no indication of the species in the dry Gori depression or in
humid Colchis, on the Black Sea coast or on the northern slopes of the
Adzhar-Imeretian ridges (Satunin, 1913; Shidlovskii, 1947, 1948, 1950).
The common vole is conspicuous on the Trialet ridge and its spurs,
e.g., the Mokrye Gory, and on Dzhavakhetia meadows.
The species is eurychoric on the Armenian Highland, where along its
lower limits the distribution increases greatly on the south and the east.
While this vole is very common on ridges surrounding Lake Sevan (Turov
and Turova-Morozova, 1928), it is also found on relatively dry and
deforested northern slopes at an altitude lower than Kedabek (1,400 m).
And similarly on the Karabakh mountain meadows (e.g., in Dali-Dag, Sary-
Yeri and Basargechar) it is widespread and yet descends to 1,200-1,300 m
on deforested eastern slopes.
On the southern slope of the Zangezur ridge in the Akulis-Chai Basin,
we found it in spring-fed marshland among heavily grazed steppelike spurs
and valleys only above 1,800 m. The common vole does not occur in the
tragacanth zone, i.e., in the ''phrygana'' and ''gariga'' formation or in the
artemisia and saltwort semidesert of the middle Araks valley and the
Kura-Araks lowland. Contrary to Ognev's opinion (1950, р. 209) and
Kuznetsov's maps (Bobrinskii, Kuznetsov and Kuzyakin, 1944), there is
no trace of this vole in the Lenkoran lowland or on Mugan. In the high
Talysh mountain it is encountered only at altitudes of 2,000-2,100 m on
steppe meadows, i.e., near the Kel'vyaz and Kosmalyan outposts. It does
not even inhabit those places inhabited by mole in the Talysh forest zone.
In general, the present-day range of the common vole in the eastern half
of the Caucasian Isthmus diverges somewhat from the distribution of the
steppe vole. In northwestern Iran and northeastern Turkey, the common
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vole is dispersed at altitudes of 2,000-2,500 m, and farther south its range
is probably higher. Blanford (1876) recorded this vole under the name of
Arvicola mystacinus De Fil. in an area between Shiraz and Esfahan,
at an altitude of approximately 2,800 m.* Goodwin recorded it from the
eastern Elburz (1939). It probably inhabits Mt. Savalan and was caught
in the region of Lake Urmia by the Caucasian Museum Expedition. In the
Georgian Museum collections there are specimens from Kars, Oltu, Great
Ararat and other localities in Turkey. The numerous vole populations
in the high Eleskirt valley in the upper reaches of the Euphrates can be
accounted for by the many Swamps and rivulets. In central Turkey some
remaining habitats are considered as relicts (Neuhauser, 1936; Ellermann,
1948).
With higher seasonal and daily temperatures, the factors of soil humidity
and mesophytic vegetation assume greater importance for the rodent in the
south, and result in an enlarged vertical distribution as the range extends
southward. They also account for the division of the range into isolated
relict islets adapted to high plateaus and ranges. The animal's range
is obviously wedged out on the high mountain ranges of the middle courses
of the Tigris and Euphrates.
The absence of the animal from western Caucasia and Transcaucasia
is adequately explained by the excessive moisture, while in the forest of
the Talysh lowland it can be attributed to dryness of the soil and vegetation.
The feeding habits of the common vole show it to be a typical herbivorous
animal. Human intervention during historical time is responsible for the
concentrated forage which comprises its present-day diet.
The main natural factor limiting this vole's population in both mountains
and plains is unfavorable climate. The influence of quadrupeds and
predatory birds on this vole during the summer season on the Caucasus
is considerably greater than on the steppe vole.
Based on its contemporary range and ecology, it is assumed that the
common vole is a European mesophilous species which developed under
conditions of moderate climate and humidity on meadow vegetation.
The explanation of the appearance and total isolation of large, common
vole populations on the ranges of southwest Asia and the Caucasus is
consonant with this concept if we presuppose an early settlement during
a humid epoch when present-day hot desert valleys and slopes were well
covered with meadow or, at the least, steppe vegetation.
From an interpretation of the contemporary ecological and distributive
characteristics of this vole, it would seem that it could hardly have appeared
on the Caucasus before the Middle Pleistocene, and then during a time of
considerable climate cooling and development of mesophytic meadow
formations in the desert and semidesert zones.
The origin of the local, high-mountain common vole in the mountains
of the eastern Mediterranean and its subsequent settlement in the north
is substantiated by the Mediterranean origin of northern mesophytic meadow
formations. Some botanists now hold the theory of an alpine origin of
meadows which were displaced to lower altitudes in the Pleistocene and
advanced in a northerly direction (Fedorov, 1952). :
Pine vole —Microtus (Pitymys) majori Thos (3. lato). Voles of
the subgenus Pitymys are known from Pleistocene deposits of Eurasia
* A vole closely related to M. guentheri Danf. et Alst. is found on Mt.Talysh and in Iran.
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and North America (Wolf, 1939; Simpson, 1945). On the Caucasus remains
of a special species of vole M. apscheronicus Arg., probably ancestral
to extant Caucasian species, are foundin Middle Pleistocene strata of the
Apsheron Peninsula. :
The distribution of pine vole on the Caucasus includes the Greater
Caucasus, the Lesser Caucasus, Asia Minor, Talysh and Elburz (Map 70).
In Ciscaucasia the pine vole can be found from the northern piedmont
forest belt up to the talus passes on the Main Range.
On the Trans-Kuban Plainthe animal inhabits some locations to an
altitude of 250-300 m above sea level, and it is common from Krymskto
Maikop on the first low ridges of relict oak forests and mesophytic meadows.
The range does not reach to the broader section of the Kuban River even
in the valley of its left tributaries — here it is replaced by the common vole.
On the east, the range enlarges and passes south to Cherkessk at an altitude
of approximately 600 m. Inthe forest zone and on the alpine meadows of the
northwestern Caucasus, the species is very common, and numerous
specimens have been collected from the Caucasian and Teberdinskii
reservations.
In central Ciscaucasia the animal inhabits the mesophytic ravine areas,
and is absent from the greater part of the dry Kabarda sloping plain.
One isolated section of the range lies in oak stands in the Pyatigor'e
area (Beshtau, Zheleznaya, Razvalka), and another more to the north
in the Stavropol uplands. In this last area the vole inhabits grazed-down
meadow glades among relict broadleaf forests, e.g., in the vicinity of
Mt. Strizhament.
The breach between this section of the range and the major one in the
Caucasian uplands is 50-60 km and is made up of steppelike foothills. There
are relict patches of pine vole habitation in forested dry ravines along the
Kalaus. On the sloping plains of the Terek and Sunzha the animal inhabits
mesophytic meadows which appeared shortly after the forests were cleared.
Contrary to Ognev's opinion (1950), this vole is found neither in the
vicinity of Mozdok nor near Grozny, where the climate is too dry for it.
In central Dagestan it inhabits meadows at 1,600-2,500 m above sea level.
In western Transcaucasia the pine vole is common everywhere from
the Black Sea coast to the mountain talus passes. In humid Colchis it is
absent only from periodically flooded alder thickets and reedgrass bogs.
The Svanetian alpine meadows are heavily populated with vole along the
fringes of birch forest grown with whortleberry and as far as the upper
sections of cereal grass—crowfoot carpets which wedge into slate talus.
Here it completely replaces the common vole. To the east, the animal is
encountered all along the southern slope of the Greater Caucasus near the
upper reaches of the Pirsagat rivulet in Azerbaidzhan.
The pine vole is sometimes replaced by the common vole in alpine
meadows in the areas of Nukha and Kutkashen. Argiropulo's statement
(1939a) that the pine vole does not inhabit the forest zone in the Greater
Caucasus is in error; on the contrary, a large pine vole population is often
observed on the forest edges and in thinned stands of oak—hornbean.
There is no indication of pine vole in the Alazan-Agrichai valley, in the
valleys of the Adzhinour Plateau or on the Kura Lowland. Claims that this
animal was found in Geok-Tepe [Geok-Chai] on desert foothills to the north
of Evlakh (Shidlovskii, 1938; Ognev, 1950) are erroneous (Vereshchagin,
1949с).
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In eastern Taurus and along the ranges of the Lesser Caucasus the animal
is spread mostly on the northern slopes and on the high humid plateaus of
Dzhavakhetia and Tsalka.
It is very common on deforested slopes of ravines of the Shakhdag Range
and in the remaining beech forests of the Terter-Chai ravines in Karabakh
where it descended to an altitude of 1,000-1,200 m. In Talysh the pine vole
inhabits only the border areas of the upper timberline, and is rare because
of the extreme aridity in summer (Vereshchagin, 1949c). The southeastern-
most section of the range is the northern slope of the Elburz Range
(Ellermann, 1948).
Pine vole is very common on motley-grass meadows near the forest's
upper edge. In these areas of ecological optimum, the greatest population
of this species can be found.
The pine vole resembles the common vole in feeding habits, but, because
of its greater digging capacity, the rhizomic constituents of herbaceous
plants feature more prominently in its diet.
The population dynamics of this vole are considerably more stable than
those of the steppe and common vole. Predatory birds inflict less population
loss on the pine vole because of its subterranean way of life and its habitat
in tall grasses and forests.
Based upon the degree of fur-cover differentiation (underfur and guard
hair), the pine vole is intermediary between the steppe and the common vole.
The pine vole is a typical mesophilous species of the eastern
Mediterranean formed on mountain meadows and in broadleaf forests. This
species is an excellent indicator species which illustrates the long existence
of mesophytic conditions on separate areas of the Caucasus-Asia Minor
landmass. The peak in pine vole population coincides with the epoch of
greatest cooling during the Pleistocene.
The Stavropol-Caucasus break in range probably occurred at the
beginning of the Holocene; the Lesser Caucasus-Asterabad break
undoubtedly happened considerably earlier.
Caucasian snow vole — Microtus (Chionomys) gud Sat. (s. lato)
(Figure 125). Fossils of this vole were found during the excavation of the
Kudaro caves in the reaches of the Rion. The animal is distributed within
limits of 1,600 to 2,000 m on the Perevalnyi and Skalistyi ridges but is
absent from the frontal chain of the Chernye Gory Mountains (Nasimovich,
1935).
Our observations in the Kuban and Teberda ravines show the vole at
1,500-1,700 m above sea level on the Skalistyi slopes and especially
numerous higher in the alpine zone on large rock taluses overgrown with
pine, birch and willow, and further on in alpine meadows near taluses and
glaciers.
At present this species is not encountered on the Stavropol uplands
(Vereshchagin and Gromov, 1953b), but it is found in the trachyte rock
taluses on the permafrost section of the Razvalka Mountain in the Pyatigor'e
area, at an altitude of 600 м. This population is now isolated from the
main range 50-60 km to the north, since the nearest vole habitat is
encountered in the canyon of the Berezovka rivulet, south of Kislovodsk
(Vereshchagin, 1953a). Higher inthe mountains, М. (Chionomys) gud
Sat. inhabits the ravines of Malka, Baksan, Chegem and Cherek, especially
rock taluses in canyons cut by these rivers in the Skalistyi ridge. It has
313
been observed in certain places in the interior longitudinal valleys at ап
altitude of 1,200-1,500m, where there are stone taluses. This vole is rare
in the Chernye Gory Mountains near Nalchik.
In the Ardon ravine the heaviest population was observed on the lateral
moraine in the pine—birch forest near the end of the Tseya glacier among
granite rocks overgrown with fern, whortleberry, sweetberry, honeysuckle
and rhododendron.
In the Terek gorges, this vole is widespread on the slope of Mount
Fetkhus near Ordzhonikidze (Formozov, 1926).
In central Dagestan numerous voles are encountered in Gunib, in the
Bogos Range and in the ravines of the Samur tributaries at altitudes of
1,300, 1,600, 2,100 and 2,500 m (Geptner and Formozov, 1941).
In August 1952, the author observed many of these voles in arid, hot
depressions in the upper reaches of the Manas brook among marl taluses
near the village of Lavashi at an altitude of approximately 1,000m. Summer
318 conditions here are pessimum for this animal.
FIGURE 125. Microtus (Chionomys) gud Sat. (s. lato)
From watercolor by K.K. Flerov
On the southern steep slope of the Greater Caucasus in eastern Trans-
caucasia this vole is found in gorges at 1,300-1,400 m, e.g., inthe
Gerdyman-Chai ravine and others, but it is common only in the cirques at
river sources bordering birch—hornbeam formations at 2,000-2,600 m. It is
also encountered on the ridges of the transverse ranges near the upper
forest belt of mountain oak at 2,200-2,700 m above sea level. Farther west
the vole appears on the Kakhetian ridge above Telavi near the snowline
(Shidlovskii, 1951). In relatively dry and deforested South Ossetia, it occurs
among the rocks from 2,000 m upward (Ognev, 1950).
314
319
In upper Svanetia this animal is very numerous in fir—maple forests.
In the arid Ingur valley it lives in sunheated gneiss rocks overgrown with
Pontic azaleas, e.g., near the village of Mestia. On steep southwestern
slopes in Abkhazia it is common from an altitude of 500 m upward to the
glaciers, although Shidlovskii (1950) recorded it for high mountains only in
the reaches of the Bzyb River. In all probability, the Surami Range is not
actually a habitat for this vole. In Asia Minor it probably inhabits the
greater part of eastern Taurus since it was found on Mount Varzambeg
south of Rize at ап altitude of 3,000 m.(Neuhduser, 1936). It was recently
observed by Shidlovskii in Dzhavakhetia as well.
The southern range of this vole is not as wide as that on the Greater
Caucasus.
Taluses of big rocks held by birch and pine forests with undergrowth of
whortleberry, honeysuckle, raspberry and Pontic azalea are densely
populatedbyM. (Chionomys).
The structure and placement of rocks and stones which ensure dry
shelters and food storage are secondary in importance for the well-being
of this animal only to the availability of the food itself. The composition
of the bedrock is less important, inasmuch as calcareous, gneiss, granite
and slate taluses are all found populated.
FIGURE 126. Reserve stocks of hay stored by Microtus (Chionomys)
Photograph by author, 1952
The composition of summer and winter foods of this vole varies greatly
according to the habitat. Nasimovich (1935) observed that in the Caucasian
Reservation the vole feeds mainly on anemones, ferns, raspberry and
Caucasian rhododendron. In the upper part of the Teberda Reservation,
Stepanov found that 32 vole stacks contained 47 plant species, of which
ferns, raspberry, mountain ash, willow, birch, cereals, strawberry,
spruce, pine and foxberry were the most frequently encountered.
315
320
According to observations made by Semenov-Tyan-Shanskii and the
author in the Tseya ravine, the animals stored mainly twigs of honeysuckle,
whortleberry, willow herb, mountain ash, Caucasian rhododendron and
stone bramble. They also feed on branches of birch and shrubs, climbing
very high on the smooth tree trunks. On the relatively dry meadows of the
Doniserdon and Baksan valleys, stacks of hay, stored in fissures in fortress
ruins (Figure 126), are composed of veronica, sage, nettle, clover, etc.
The Caucasian vole is adapted to rock reliefs and avoids digging. Among
the other voles, it stands out in its astonishing climbing ability, almost
the equal of that of the dormouse or the squirrel.
Notwithstanding the fact that great variations in the population of this species
have not been noted, in certain years and in certain gorges they are rarely
encountered. This may indicate the incidence of local epizootic diseases.
This species is not as vulnerable to climatic affects as are other vole
species.
Of the predators, only marten, weasel and large owls are dangerous
to this vole, although any newly-settledcarnivore, like the sable, may prey
upon it.
Judging by the large population it supports and the size of the specimens
it produces, the subalpine zone of the western part of the Greater Caucasus
provides the optimal contemporary ecology for M. (Chionomys) gud.
The formation of rocks during orogeny and glaciation was often more
important to the development of this vole's range than the presence of
mesophytic vegetation, as can be seen by the densely populated, dry
interior valleys and by central Dagestan where the ecology of this species
shows features similar to those of Microtus (Chionomys)
nivalis Mart. (s. lato). An analysis of the range and habitat brings us
to the conclusion that the Caucasian vole is an early local species,
associated in its evolution with the development of high-mountain plant
formations of the Miocene. The glaciation of the Greater Caucasus in the
Pleistocene could not have displaced the populations of these animals to the
plain, as thought by some faunists and paleontologists; it only partially
lowered their range. Even during the lowering of the snowline to 1,000-
1,100 m, a wide habitat for vole existed on the meridian ranges and onthe
frontal longitudinal ranges. However, the cold and humid climate during
the glaciation, whichbrought about the development of meadow formations
in the foothill plains, created the conditions for the penetration of the
animals into places below river bank escarpments. The vole probably
appeared in the Pyatigor'e area during one of the cold epochs, but it is
doubtful that it reached the Stavropol uplands. Future fossil finds in the
foremontane regions from the Middle Quaternary may fix this history more
precisely.
Nowadays, cattle grazing and the felling of mountain forests diminish
the range of the Caucasian vole in the eastern Caucasus, but enlarge it in
the western Caucasus.
Asia Minor snow vole —Microtus (Chionomys) nivalis Mart.
(s. lato). Remains of this species were recorded from Pleistocene and
Holocene strata of many caves in England, France, Germany, Italy, Austria,
Sweden, Czechoslovakia and Hungary (Wolf, 1939).
The remains of a special species —M. (Chionomys) machintoni
Bate — found in Acheulean to Mesolithic deposits (Bate, 1937) are recorded
1704 316
for the caves of Mount Carmel in Israel. In the U.S.S.R. only remains of
Holocene Age are known on the Armenian Highland. The contemporary
species is widespread in Mediterranean mountains — from the Pyrenees
to the Balkans, in Syria, Palestine and Asia Minor, onthe Caucasus andon
the western Kopet Dag. In the upper Pleistocene this vole probably inhabited
even the Balkan Mountains.
On the Greater Caucasus, the animal is known only from the northern
slope of the range: the Fisht, Pshekish, Bolshoi Tkhach, Acheshbok and
Pambak peaks, and from the upper Teberda and Cherek river gorges; it is
not encountered in the Balkarian Cherek or farther east. The paleontological
sites on the Greater Caucasus are located at altitudes of 1,500-2,000 m in
the subalpine zone. The habitats here are the same as those of the Caucasian
_ vole (Map 67).
321 On the Lesser Caucasus the Asia Minor vole inhabits the forest and
alpine zones on the Trialet ridge, in Dzhavakhetia, on all ridges of the
Armenian Highland, in Karabakh and in Talysh.
The vertical distribution increases in a southeasterly direction according
to the locations of phytolandscape zones, although there are exceptions.
On the Trialet ridge the species has been found at altitudes of 2,000 m near
Bakuryan; on the Pambak ridge it is common at altitudes of 1,800-2,500 m;
and on the Shakhdag it lives in the Shamkor-Chai ravine between 1,600 and
2,300 m. Southward, it is encountered in the xerophytic mountain steppe,
e.g., near Sevan on the Saraibulakh and Daralagez ridges (Turov and
Turova-Morozova, 1928; Dal', 1944b, 1949b). Above Yerevan it occurs at
a height of 1,200 m in rock taluses covered with shrubs (Flerov and Gureev,
1934).
Along the Akulis-Chai ravine on the southern slope of the Zangezur, in
1947, we foundthe Asia Minor vole at altitudes from 1,600 to 2,500 м in
rock taluses overgrown with spiraea, dog rose, almond, hawthorn and
buckthorn in the zone of astragali—traganth.
It was not encountered by the author in the beech forest in the Terter-
Chai gorge in Karabakh, but is found higher in the alpine zone near
Kelbadzhar.
In Talysh and Kopet-Dag, there is only a relict range. The author found
a small vole population in June 1945 on the bald Kelakhan Mountain in the
dry Diabar depression at 1,900 m. The animals were encountered here in
the taluses of teschenite rocks with a sparse xerophytic cover of saxifrage,
oxeye daisy, brome shoots and xerophilic ferns among the mountain-
saltwort semidesert (Vereshchagin, 1945b). This vole probably inhabits
Savalan and the Elburz Range.
The Asia Minor vole achieves its maximum population in a zone of steppe
meadows, e.g., near the Sevan shore. The rocky substrate is required
for the well-being of this species. The habit of storing food is not so
strongly developed as it is in the Caucasian vole, but the capacity to adjust
its habitats to varying conditions is immeasurably greater; it adjusts to a
wide range of feeds and of humidity. Whereas in the subalpine zone of the
western Caucasus this vole lives side by side with the mole and the earth-
worm, in Talysh and on the southern slopes of the Armenian Highland it is
found in the mountain xerophytic zone living with the tarantula and the
scorpion. Onthe whole, the vole is more adapted to low pressures and
rocky substrates and less to succulent foods and low temperatures.
317
The diminution in the animal's size in its eastern range and its
disappearance from the medium-altitude southern slopes of the Armenian
Highland indicate that the southeastern sections of the range are already
located in a contemporary pessimum region.
Discontinuities in the southeastern areas of this vole's range and the
isolation of the mountain habitats in Talysh and Kopet Dag are explained
by the increasing aridity in the eastern Mediterranean which followed
optimal conditions for the settlement of the species during a pluvial cold
epoch.
However separated and various the isolated ranges in the Mediterranean,
they indicate that the snow voles of the Pyrenees, Alps, Carpathians and
Caucasus are separate species of parallel development from genetically
related material.
322 The postglacial rise in temperature and the elevation of phytolandscape
zones have reduced the vole range, deepening it and isolating different
geographic races of the snow vole.
The limited spread of the snow vole on the Greater Caucasus is partially
explained by the insufficient adaptation of this southern species to the
marked humidity of the southwestern ridges; it is more difficult to explain
its absence from the dry longitudinal valleys of the northern slopes of the
eastern ridges and from Dagestan.
Long-tailed snow vole — Microtus (Chionomys) roberti Thos.
A unique finding of Quaternary fossils is recorded from the Upper Paleolithic
of the Sakazhia cave in western Transcaucasia. The contemporary
distribution of this vole includes the northern slopes of eastern Taurus in
Asia Minor, the Adzhar-Imeretian Range, the Trialet ridge and the Greater
Caucasus (Map 69).
In the northwestern Caucasus this vole is found on the Chernye Gory
Mountains from elevations of 800-1,000 m to the alpine zone. It is rare in
central Ciscaucasian gorges and is not encountered at all on the Stavropol
uplands. It is common in North Ossetia at altitudes of 800-1,000 m.
In Dagestan it inhabits beech forests in the upper reaches of Andi Koisu
near the villages of Takhota and Choroda, but it is probably absent from
other parts of Dagestan (Geptner and Formozoy, 1941), as it still is from
the inner longitudinal valleys of the northern Caucasus.
The author found the vole in Azerbaidzhan (Vereshchagin, 1940a) in
1935 in the Katekh-Chai ravine, northwest of Zakataly, at an altitude of
1,450 m, butitis not known farther east.
The vertical distribution of M. (Chionomys) roberti Thos. on the
BlackSea slope of the Greater Caucasus is from sea level to the upper
forest belt (Shidlovskii, 1947, 1950). In Adzharia the animal inhabits the
forest zone from 800 to 1,800 m (Shidlovskii, 1948). It is distributed in all
of South Ossetia except the dry Gori depression.
A typical well-populated habitat of this vole is a shady, humid, dead-end
gorge. In the western part of the Greater Caucasus and on the Lesser
Caucasus, such biotopes are to be found in fir and fir—beech forests with
tall broadleaf grasses of burdock, cow-parsnip, nettle and orchard grass
covering the bottoms of creek valleys. In Ciscaucasia the population is -more
numerous in the lower humid area of the Chernye Gory Mountains than in
the upper parts of large ravines. In eastern Transcaucasia the opposite is
observed.
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323
According to its morphological type, particularly its fur cover, this
vole resembles the water vole to a certain degree. The animal often swims
in mountain brooks and in general lives in conditions of extreme humidity.
The speciation of this vole is associated with the early mesophytic and
even hygrophytic phytolandscapes and biotopes which existed in the
mountains bordering the Black Sea depression on the southeast and east
in the Miocene. The reduction of the M. (C hionomys) roberti range
to the east completely corresponds to the changes in landscape and biotopes
in the mountain forest.
The existence of an isolated, actually a relict, population in the upper
reaches of the Andi Koisu in Dagestan can be explained by the penetration
of the animal from the southern slope during a humid epoch. Tumadzhanov
(1940) recorded a similar immigration of the beech into Dagestan in the
Holocene.
In all, М. (Chionomys) roberti is an endemic indicator species of
the ancient mesophilous faunal complex in the Caucasus.
Common red-backed vole —Clethrionomys glareolus Schreb.
Fossils belonging to the genus are known in Europe from the Pleistocene.
The main part of this species' range is located in taiga and broadleaf forests
of Eurasia. The animal penetrates into the steppe through the forested
areas of river valleys. Ecologically this vole is mesophilous and
psychrophilic. Only the Pontic subspecies, C glareolus ponticus
Thos., is found in the southeastern Mediterranean, inhabiting the northern
slopes of Taurus on the southern coast of the Black Sea (Thomas, 1906;
Neuhauser, 1936) and the Adzhar-Imeretian Range (Map 72).
In Adzharia the animal is found in spruce forests along the Supsa River
(Shidlovskii, 1940a). This southern section of a wide range, now isolated
by the Bosphorus, is a remarkable example of the southern faunistic
influence on the Caucasus through the Balkans and Asia Minor.
The common red-backed vole's absence from the western part of the
Greater Caucasus indicates that it penetrated the southern Black Sea coast
very late, during the period of maximum cooling in the Upper Pleistocene.
Steppe lemming — Lagurus lagurus Pall. Fossils of this Upper
Pleistocene species are known from several localities on the Russian
Plain — the Don, Volga and Ural valleys. The present range is located in
the steppe and semidesert zones from the lower reaches of the Dnieper
to the upper reaches of the Yenisei and the borders of northwestern China
(Vinogradov and Argiropulo, 1941). The steppe lemming is also widespread
оп areas adjacent to the Caucasian Isthmus, on the northern coastlands of
the BlackSea, the Sea of Azov andthe CaspianSea. Onthe Isthmus it is found only
in the northern Ciscaucasian steppes in the interfluve of the Kuma, Kalaus
and Manych, i.e., northwest of the Stavropol Plateau (Sviridenko, 1928).
Kistyakovskii (1935) found remains of this animal in bird pellets in the
vicinity of Achikulak, Blagodatnoe, Sukhaya, Padina, Petrovskoe and
Kambulat (Map 64).
The steppe lemming appears in Ciscaucasia only in peak breeding periods
or during migrations provoked by unfavorable conditions. We assume that the
steppe lemming penetrated only recently into the Caucasus Isthmus, probably in
the Upper Pleistocene or Holocene, from the adjacent steppes of the Russian
Plain, drawing the inference from its underdeveloped range in the
Ciscaucasian steppes, from its stable connections with the landscape, and
319
from formations of cereal grass—motley grass, feather grass—fescue and
steppe—wormwood of the temperate belt of Eurasia. This settlement could
have taken place at the time when the Manych strait disappeared.
Promethean vole —Prometheomys schaposchnikovi Sat.
(Figure 127). Fossils of this vole carried in by the eagle owl are known
from Upper Paleolithic strata in the Gvardzhilas cave in the Kvirila River
ravine in eastern Transcaucasia at an approximate altitude of 450-500 m,
and from Lower Paleolithic strata of the Kudaro I cave.
The Gvardzhilas cave is located outside the contemporary range of this
animal, but to study the shifts in range during the Upper Pleistocene, it is
necessary to follow the features of this vole's contemporary distribution
on the Rachin ridge and to investigate the radius of the eagle owl's hunting
324 flights in this region.
FIGURE 127. Promethean vole
Watercolor by E. Ya. Zakharov
320
325
The present distribution of Promethean vole is confined to the western
half of the Greater Caucasus and to the Adzhar-Imeretian Range of the
Lesser Caucasus (Map 65). On the Greater Caucasus it is encountered
from the Fisht and Oshten peaks to the Kazbek-Krestovyi Pass and Gudaur
Dag. It has been traced in the Caucasian Reservationin the Mzymta valley,
along the lower Kardyvach Lake, in the meadows where the Kurdzhips and
Pshekish rivers rise, in the Kisha and Urushten basins, in the Malaya
Laba sources, and near Krasnaya Polyana (Turov, 1926a, 1926b). The
animal was caught in Abkhazia at the Avatkhar site at an approximate
altitude of 1,500 m. In Svanetia in 1948 the author followed the animal on
high-mountain meadows in the upper reaches of the Ingur and the Rion and
found it particularly numerous on the Zagar pass of the Lechkhumi ridge
at altitudes of 1800-2,800 м. This rodent is not encountered in the dry
longitudinal Ingur valley. In South Ossetia the Promethean vole was hunted
in the Bolshaya Liakhva upper reaches (Shidlovskii, 1951). The animal
was traced on the mountain meadows of the Mamisson pass to the railroad
stations Kazbek and Gudaur in the central part of the Greater Range (Turov,
1926a, 1926b). The author found neither the animal nor its tracks in the
upper reaches of the Balkar Cherek, Doniserdon and Ardon. We have no
data on the presence of this vole in the upper reaches of the Kuban, Baksan
and Malka. Thus, the range on the northern slope of the Main Range
is broken at least from the Laba basin and Zelenchuk to Ardon. This may
be explained to some extent by the destructivity of mountain glaciers.
On the meadows of the Skalistyi ridge in Ossetia, in Kabarda and in the
Grozny Region, this rodent was not observed in places perfectly adapted
for its habitation. Apparently, the Promethean vole was not distributed
north into the dry longitudinal valleys even during the Pleistocene.
Nor has it been observed east of Kazbek on the Perevalnyi Range. Inthe
Zakataly Reservation, which the author has carefully explored, it is absent,
as it is from central Dagestan. The rapid thinning out of the rodent
population east of the Caucasus can be attributed to the great reduction in
area of alpine meadows and to xerophytization as shown by the development
of sheep's fescue and matgrass, instead of a motley-grass community.
On the Lesser Caucasus, the Promethean vole has been found only in
the upper reaches of the Bakvis-Tskali River, the left tributary of the Supsa,
near the Bakhmaro health resort, at an altitude of 1,600 m (Shidlovskii,
1940a, 1947); these subalpine meadows are located west of the Gotimaria
(Metis-Tskaro) peak (2,868 m). The range in this area is considerably
wider; according to Shidlovskii's data, the vertical distribution lies within
limits of 1,500-2,800 m, but the animal can also be found at lower altitudes.
The contemporary ecological optimum of this animal is within subalpine
and alpine meadow zones with a predominance of lady's mantle and bunch
grasses (orchard grass, meadow grass and others). Meadows lying among
thickets of Caucasian rhododendron and birch are also densely populated.
The rodent does not enter the rhododendron growth where food is not
available, but in birch forests and on the upper forest edge grown with
tall grass it is as common as it is on open meadows. This vole's habitation
of beech forests in the Caucasian Reservation, noted by Turov (1926b),
may be, in some cases, the consequence of recently advancing forest
formations into the alpine meadow zone. In heavily shaded beech and
hornbeam forests the vole is never found because of the lack of grass food.
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326
In many places, the animal inhabits oat, wheat and potato fields or invades
their borders, as in the upper reaches of the Ingur.
Its strictly subterranean way of life and its diet of grass roots are
reflected in the animal's morphology: flattened skull, extended incisor
curve, elongated diastema, rudimentary eyes and shortened extremities.
The skull, witha crest extending nearly to the occiput, is very similar to
that of the muskrat. In its weakly-differentiated and vertically-disposed
fur cover and coloration (often black), the animal is similar tothe mole vole.
In general, in a morphological-functional sense, the Promethean vole
combines features of the muskrat, water vole and mole vole, but, in
contrast to the latter, it lives in mesophytic conditions (see also Vinogradov,
1926; Ognev, 1926b). According to Gambaryan's observations (1952), the
Promethean vole, unlike the mole vole, combines terrestrial feeding with
root grubbing.
Such adaptations indicate the early origin of the species and its pristine
associations with the landscape of the high-mountain mesophytic meadows.
The population fluctuation of the Promethean vole can be attributed to
mass death during the spring showers and after frosts when the water
freezes in the burrows (Turov, 1926a).
The strict endemism of this species shows the conservation of the
ecological conditions on the western Caucasus: the ancient origin and
continuous existence of the landforms and the zone of mesophytic subalpine
and alpine meadows from the Lower Pliocene at the latest.
The division of the range into two parts — the Caucasian and Lesser
Caucasian — may have taken place by postglacial time as a result of the
drying-out of the meadows on the Surami crystalline rock massif.
This vole is an early Pliocene species endemic to the Lesser and
Greater Caucasus with a relict range which has increased somewhat in
present time because of the expansion of the high-mountain-meadow zone
through forest-felling and animal-grazing.
Water vole — Arvicola terrestris L. ($. lato). Fossils of this
vole are known from Pleistocene strata in Europe.
Remains of Arvicola terrestris L. (s. lato) are common on the
Russian Plain in Upper Pleistocene and Holocene deposits. South of the
Caucasus the remains of this vole have been found in Acheulean strata of
Syrian caves. On the Caucasian Isthmus there have been isolated fossil
finds in Middle Quaternary strata of eastern Transcaucasia and inthe Upper
Paleolithic of western Transcaucasia, and numerous finds in Holocene
deposits in caves which are day rests for eagle owls.
The contemporary distribution of the species includes the forest, steppe
and semidesert zones of Eurasia. South of the Caucasus it can be traced
in northern Iran and Asia Minor as far as Palestine.
On the Caucasus the water vole inhabits small areas from the level of
the Caspian to an altitude of 2,600-2,800 m, but is not found in broadleaf
mountain and coniferous forests, or in rocks and taluses of passes (Map 66).
Its greatest population is observed on the floodplain meadows of the lower
Don and a somewhat smailer population in the Kuban floodplain.
In the Colchis and Lenkoran lowlands it is very rare, although it is found
in damp depressions which flood annually. In the eastern Ciscaucasian
semidesert and in Transcaucasia, it is common on lakes and streams
overgrown with reed and rush, where it lives in the summer months and
builds sphericalnests among the thickets, but does not approach the shore area.
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327
The largest population of water vole in the foothills is observed on spring
rivers like those on the Terek-Sunzha Plain.
In the interior steppe valleys of the Greater Caucasus northern slopes
and in central Dagestan, a peculiar type of vole, ecologically distinguished
by its subterranean existence, is found. In these areas this vole inhabits
the bottoms of gorges, particularly irrigated farmland. In the alpine zone
of the Greater Caucasus it occurs rarely and only in the upper reaches of
the Terek and the Aragva.
On the Armenian Highland, in Asia Minor and in Iran, the water vole
is common on subalpine meadows on the upper reaches of rivers (e.g.,
in Karabakh, in the upper reaches of Akstafa and in the lake region of
Dzhavakhetia). In the Araks valley many of these voles are found near the
spring rivers of the Aralik depression.
It can be seen that this vole is widely distributed and especially attracted
to water biotopes.
The ecological optimum of the subterranean populations of these voles
is at mean altitudes in the dry area of Caucasian plateaus; that of the
hydrophilous populations is in the semidesert zones which lack excessive
atmospheric humidity[sic!]. The low population density of the water vole
on the Black Sea coast and in the Colchis and Asterabad swamps indicates
a certain degree of similarity to the distribution of the common vole.
In the Colchis swamps it is the tremendous competition and persecution
from the Norway rat that produces this reduction in population, and in
Asterabad the drying-up of reservoirs in summer and the compaction of
the Lenkoran yellow podzolhave the same deleterious effect (Vereshchagin,
1941b, 1949d).
Subterranean mountain populations differ from hydrophilic species in
elongated diastema, greater curvature radius of the upper incisors, and
shorter tail length, and the divergence in the contemporary ranges of the
respective Species can be observed along a line of secondary remove from
the water medium and from these distinctions in features.
In southwest Asia and onthe Caucasus the formation ofthese characteristics
is probably related to the uplift of mountain systems during the Quaternary.
Paleoecological data suggest that the water vole ofthe Caucasus is probably
of Upper Pliocene origin. In Transcaucasia the southern, relatively
xerophilous populations actually predominate.
The consistency of the species range in the Mediterranean is ensured
by both the natural and artificial development of freshwater biotopes and
the ecologic plasticity of the animal.
Order PERISSODACTYLA
Representatives of the families Equidae and Rhinocerotidae appear in the
Quaternary fauna of the Caucasus, but by the Holocene only Equidae remained.
The historical origin and disappearance of this family is examined below.
Family EQUIDAE
The earliest fossils of the large, true horse, Equus stenonis Cocchi,
are known from the Caucasus and adjacent territories from Upper Pliocene
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deposits. Probably the horse appeared simultaneously with the southern
elephant introducing a new phase in the fauna. Later, in the Middle
Pleistocene, the ''semi-ass'’ appeared on the Caucasian Isthmus.
The history of true horses can now be traced on the Caucasus very
schematically from a number of deposits of various ages.
Quaternary horse — Equus caballus ($. lato). After Equus
stenonis, remains of which were found in Akchagyl and Apsheron deposits
of the Isthmus, the Taman Upper and Lower Pliocene horse, E. aff.
sussenbornensis Wist., found in conglomerates of the Taman
Peninsula and the ancient Caspian terrace shingle near Baku, is the closest
to the Quaternary horses (Chapter II and Map 79).
Fossils of Midde Pleistocene horses (which are not recorded in detail)
are abundant in Binagady asphalts and are found isolated in clayey beds of
the Khazar stage in many localities in the Stavropol area and the Krasnodar
Territory. These horses were related to Е. caballus chosaricus
Grom., found in Middle Pleistocene deposits of the Russian Plain.
Remains of horses from the Il'skaya Paleolithic settlement in Ciscaucasia
and from the Paleolithic sites of Imeretia and the Lesser Caucasus uplands
are probably related to Е. caballus latipes Grom. of the Russian
Plain (see Gromova, 1949). South of the Caucasus, remains of Upper
Pleistocene horses are known from Paleolithic strata in the Tamtama and
Bisotun caves, from talus loams in the Maragheh vicinity of Iran and,
southwest, from Palestinian and Syrian caves. The southern horses were
of a lighter build than the northern animals.
Specimens of Pleistocene horse have been taken in the following
proportions to other species from sites representing various types of burial:
6.6 % from the Il'skaya encampment; 5.2% from the Binagady asphalt;
2.5% from the Sakazhia cave; 16.5% from the Mgvimevi shelters; 50.0%
from the Zurtaketi encampment.
Assuming that the forms ofhorse we have mentioned, which inhabited the
Isthmus during the Pleistocene, constitute a unique genetic series, then
the origin of the Caucasian Holocene horse should not be sought in migrated
forms originating elsewhere.
It is difficult to separate the remains of wild horses which lived on the
Caucasus in postglacial time from the remains of domesticated horses.
In the strata of post-Paleolithic encampments, it is quite common to
find bones of horse among kitchen middens and skulls and entire skeletons
in the deposits. It is very probable that most of these remains belong to
tarpans —Е. caballus gmelini Ant.—or to their domesticated offspring.
The records of horse remains found on the Caucasus in kitchen middens
of encampments and in deposits fromthe historical age show that this animal
had an enormous economic, dietary and ritualistic importance for various
tribes and peoples (Table 71).
Numerous remains of horse are known also from Urartu burials of the
10th to 12th centuries A.D. in the Sevan Basin (Lalayan, 1929) and from
the cultural strata of the Later Bronze Age and the Middle Ages near
Samtavro and near Mingechaur.* The horse (including the wild horse)
was among the preferred subjects of Bronze Age drawings and sculpture
in Southwest Asia and in the Caucasus.
In the middle of the first millennium B.C., wildhorses stillinhabited
the Caucasus in large numbers. These were thin-legged animals with little
* Materials from these localities have not been preserved.
324
ears оп small heads similar to the Arab horses.
Their slight build and
refined form had nothing in common with the body type of tarpans.
Marvellous reliefs on the walls of the Assyrian palace in Nineveh (Atlas
of Ancient History of Egypt, Southwest Asia, India and China, 1937, Table
156, Figure 2) show hunts in which they were set upon by dogs and brought
down by spears.
This accumulation of evidence confirms the fact that the
tarpan was an aboriginal horse of Europe and the northern half of Asia and not
a settler from the south at the end of the Pleistocene (much less during the
Holocene) as is held by some paleontologists.
In the monuments of the Koban cultures in Ossetia and Abkhazia the
horse was depicted on belt buckles and ax handles as a rather slender and
long-tailed animal. Bronze figurines of horses with heavy manes and long
tails are among the material obtained by Uvarova (1900) in Ossetia.
Without taking into account the stylization of the representations, one sees a
similarity in body frame, particularly in the light head of this animal,
to the southern type of horse, in marked contradistinction to the tarpan.
strata of encampments and burials on the Caucasus*
(329 ) TABLE 71. Number of kitchen and ceremonial remains of horse, compared with other animals, in
Cece ess a a
Regions and localities
Dating
P
roportion of horse
specimens to
other specimens
(in %)
Ciscaucasia and Russian Plain
Sarkel fortress (Slav and Khazar strata) ........ 8-18th centuries A.D. 10.3
Settlement near Tsimlyanskaya (Khazar strata) ... 8-10th centuries A.D. 37
Settelement of Zeyukovo near Nal'chik ........ 6-8th centuries A.D. 16.7
Ancient town sites: Isti-Su and Alkhan-Kala in the
Sumanay Walley oe в 0155 ows 6 aha So diss ae 616 2-3rd centuries A.D. Toll
Burials near Ust-Labinskaya ...........+..-.- 6th century B.C. — 2nd
century A.D. 21.2
Site of ancient town of Alkhaste on the Assa...... 6-5th centuries B.C. 13.0
Site of ancient town of Semibratnoe on the Kuban ..| dth-1st centuries B.C. 17.0
Cepi 5
Ancient Greek t f the 4
ee И Я Phanagoria 5th- Ist centuries B.C. 12.3
| Taman [Hermonassa ] 14,1
smell Cm wos MEM hs бо мо бо poo mecds eo Oc Ist millennium B.C. 12.5
mud-hut 1 Second half of 2nd 132
Tsimlyanskaya on the Don: {
у у mud-hut 2 millennium B.C. 12.0
Eastern Transcaucasia
CGhuakhuraKabalasiortressiew. bares. ое ens 12-14th centuries A.D. 16.6
Settlement in the region of Baku fortress ....... 9- 12th centuries A.D. 8.1
Settlement пеаг Sumgait и. 266 ee ee ee 1st millennium B.C, 8.3
Western Transcaucasia
Akhshtyrskaya cave near АЧ1ег.............
* Author's unpublished data,
Neolithic and later strata
325
329
330
331
In large Scythian mounds in Ciscaucasia and south of the Russian Plain,
archaeologists often encountered mass ritual burials of horses composed
of some tens and even hundreds of specimens (as many as 300).
In Scythian memorials realistic representations of horses are quite
common. A drawing of a stallion with a short-standing mane, thick legs,
heavy head, small ears and blunt muzzle resembling а tarpan is depicted
on a silver dish from the Maikop burial (Farmakovskii, 1914; Figure 128).
The well-known horse representations on the silver vase from the Kul-Oba
burial probably portray domesticated horses. Outline drawings of horses
are also found on the rocks of Kabristan, south of Baku. These were made
probably during the time of the ancient Romans or earlier when wild horses
lived on the Kura steppes (Vereshchagin and Burchak-Abramovich, 1948).
Pallas, Gmelin and Guldenstaedt reported on tarpan in southern Russia
and on the Ciscaucasian steppes at the end of the 18th century . Levchenko
(1882), Keppen (1896), Anuchin (1896), Geptner (1934b), Antonius (1938)
and others published many articles and notes tracing the history of the
disappearance of this animal in the second half of the 19th century. A
particularly complete and impartial survey on the investigation of the tarpan
was submitted by Keppen, who insisted on the necessity for studying horse
skeletons from the southern Russian burials. This investigation has not
been carried out however. *
Unlike most investigators, Anuchin did not consider the wild horse of
the Eurasian steppes and deserts as a species, but thought it to be a type
of horse which belonged to nomads and ran wild. It is obvious that, if this
were the case, these horses ran wild on a large scale, and such horses
must have closely resembled the original form — the wild horse of the Upper
Paleolithic and the Lower Neolithic. There is little data on the tarpans
in the Caucasus. Rovinskii (1809, р. 131) wrote: ''Wild horses or tarpans
are found in flocks in the Kuban steppe, near the Ural mouth and farther."'
The author could find no historical data on the presence of wild horses
in Transcaucasia. By the Middle Ages, the only survivor of the dense
early population of wild horse on the northern plateaus of southwest Asia
was the kulan, since the horses had probably been caught and domesticated
by Hittites, Assyrians and Urartus long before the Christian era.
The rapid evolution of horses in the Quaternary is usually associated
with the subsequent development of steppe topography and coarse grasses,
some of which — the grain formations — appeared as early as the Pliocene.
The adaptive features and contemporary ecology of wild and
domesticated horses corroborate this association. Notwithstanding this
typical association, horses could still subsist on the forest steppe and even
in the forest zone of Europe because, in large herds, they can actively
defend themselves against packs of predators (e.g., wolves), and because
they can grab for food under the snow cover. In these areas, they fed in
glades and swamps just as the extant domesticated horses of Yakutia do.
The characterization of the Quaternary horse as a stenotopic steppe
animal did not do justice to its conformation, which represents a universal
form in ungulates. This is substantiated by the success in using domestic
horses in widely differing topographies — from the glaciers of Greenland,
Antarctica and the Alps to the tropical forests, swamps and deserts of all
continents.
* Skulls and skeletons of horses from earlier diggings in large burials in the Ukraine and Ciscaucasia have not
been kept.
326
(330)
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FIGURE 128. Representation of a horse, primitive bulls, lions and other animals on a silver vessel
from Maikop hill (Farmakovskii, 1914)
Recently Vetulyani (1952) developed Keppen's theory that, especially
in Middle Europe, wild horses were forest animals. As Vetulyani saw it,
the tarpan adapted itself to a middle-forest-belt habitat and, in Europe,
became a forest animal only during the last stage of its development, иен
in the historical epoch.
New observations on artiodactyls would indicate that such a change in
biotopes resulted from the development of forests in former forest-steppe
areas and the hunting down of horses by man.
The intersecting relief and relatively dense population of the Caucasus
did not favor such adaptations. The steep slopes, both afforested and
deforested, were unsuitable for horses, and the gentle slopes were toc
exposed.
The date of the disappearance of the tarpan from the Russian Plain
has been previously established as the 1870's, but Geptner's recently
published data (1934b, p. 482; 1955) on the last tarpan to inhabit the Poltava
Region considerably advances this date to 1918-1920.
Tarpan extinction on the Caucasus resulted from their harassment by
nomadic horsemen, steppe plowing, human settlement near watering places
and, finally, from hunting with firearms (see Keppen, 1896).
lato). The fossil remains of
Kulan —Equus hemionus Pall. ($
Equus hemionus Pall. are known from Pleistocene and Holocene strata
of Eurasia, particularly in steppe and semidesert zones. In Gromova's
opinion (1949) kulan fossils recorded from Western Europe as
327
Е. hemionus fossilis Nehr. may be related to small horses. However,
the possibility of an early penetration of kulan into the west from Asia
during the postglacial transformation of plains into steppes cannot be
excluded. Phalanges and metapodia of semi-asses from upper layers of the
Afontova Gora [Krasnoyarsk area] on the Yenisei, from sand dunes near
Troitskosavsk [now Kyakhta] on the Selenga, from Irtysh and from Altai
caves (Gromov, 1948; Vereshchagin, 1956) are mainly from the Upper
Pleistocene and Holocene. Fossils from the Upper Pleistocene of China
have also been recorded.
In central Asia remains of kulan are known from Paleolithic d2posits
in the Bukhtarma cave (Vereshchagin, 1956) and from Bronze Age
strata in the Kaunchi-Tepe cave [Tashkent area]. Duerst (1908) recorded
the remains of a thin-legged horse, Е. caballus pumpellii Duerst,
fromthe earliest Annau strata (3000-1500 B.C.) near Ashkhabad. Gromova
(1949) is of the opinion that these animals may be kulans.
Kulan lived south of the Caucasus from the Lower Paleolithic Acheulean
up to the present (Bate, 1937). Kulan fossils are also recorded from
Paleolithic strata of the Barda-Balka cave in south Kurdistan.
On the basis of contemporary data, the wild ass (Figure 129) is
undoubtedly a southern species which originated in the Eurasian semidesert
and upland steppe.
The vague relationship of the Pleistocene ass, Е. hidruntinus Reg.,
with the wild ass makes it difficult to establish when and by which routes
the wild ass emerged on the Caucasian Isthmus. Up to now a direct
phylogenetic link between these species was generally denied on the grounds
that the wild ass had a more primitive dentition (Stehlin and Graziosi, 1935;
Gromova, 1949). If we consider as an analogy related orders of Quaternary
elephant, rhinoceros, horse and primitive bison, it is possible to conclude
that the wild ass is a direct descendant of the Pleistocene ass. The claim
of an autochthonous origin for the wild ass on the Caucasus follows from
this conclusion. Remains of Holocene wild ass are known from several
localities on the Caucasian Isthmus and in adjacent areas of the Russian
Plain (Map 78). The oldest finds are from a settlement of the later half of
the second millennium B.C. near Dzhemikent, south of Makhachkala; the
latest finds are from 13th-century strata of a settlement in the vicinity
of Baku bay.
Historical data on the wild ass on the Caucasus and in adjacent territories
are fairly numerous. Xenophon (1896 edition) gives a vivid description of
ancient Greek wild ass hunts; these wild asses lived together with ostriches
in the wormwood steppes of the upper reaches of the Euphrates. The records
of Moisei Khorenskii (1893 edition) state that Armenian kings of the
Arsacid Dynasty (1st-2nd century A.D.) hunted onager (wild ass) and boar
in the middle Araks valley. These records were recently discussed by
Sarkisov (1946a) who mentioned that onagers had been recorded in Armenia
in the ''Code of Laws of the Armenian-Gregorian Church"' in the Geography
of Armenia by Ananiah of Shirak(7th century) and by Grigor Narekatsi
(11th century).
Under the influence of Satunin's ideas, Sarkisov assumed that the
mountain barriers eliminated all penetration routes into Armenia for the
onager except one from the ''Aral-Caspian lowland. '' Clearly, mountains
are no barrier to this species, since, in historical time, onagers inhabited
328
every intermontane valley that was even slightly accessible, including the
uplands of southwest Asia.
Even more convincing are the arguments to substantiate the existence
of the wild ass in Armenia in the book of mathematics by Ananiah of Shirak
which is discussed by Ter-Pogosyan (1947).
The Armenian historian of the 10th century Moisei Kalankatuiskii
(1861 edition) mentioned the wild asses inhabiting the Barda kingdom
(in the contemporary Mil'skaya-Karabakh steppe). Nizami Ganjawi
(12 century) praises the wild ass hunt in the Kura-Araks lowland. Rashid
ad-Din (14th century, 1946 edition) mentioned that among the victims of the
grand hunt organized by Gazan Khan in Talysh were wild asses. At the
beginning ofthe 17th century Adam Olearius (1870 edition) recorded an
onager hunt by the Iranian shah on onagers or Persian ''gurs'' in one of the
sections of an enormous reservation in the vicinity of Iran's capital,
Esfahan.
More recent data on the wild ass developed by Russian naturalists
indicates that these animals inhabited only the area beyond the Volga during
the time of Pallas.
Levanevskii (1894) gives a good description of the habits of this species
in the region of the Kirghiz steppes of the Emba District. From this
description it seems that the wild ass was common in the semidesert
between Emba and the Urals even in the second half of the last century.
333 It was probably at this time that the wild ass population declined rapidly
and the range shifted to the east and southeast. It is difficult to relate this
shift and reduction in population only to the extermination of the animal
with improved firearms or to an increase in human population. The Nogai
and Kazakh retainedtheir mainweapons, percussion muskets and flintlocks,
from earlier times. The increase of encampments was limited by the fodder
capacity of desert and semidesert pastures. It is therefore probable that
other causes influenced the dynamics of the animal population.
By the beginning of this century the wild ass was found in the U.S.5.R.
only in the semideserts of northeastern Kazakhstan — Bet-Pak-Dala and
Semirech'e. These animals disappeared from these areas and from
southern Turkmenistan-Badchis in the 1930's. The peculiarities of wild
ass distribution in the Iranian uplands and in Arabia are as yet unknown.
Taking into account the stenotopic character of the wild ass and its
adaptation to desert-steppe topography, one can fix its appearance on the
Caucasian Isthmus at the beginning of the xerothermic period. It may have
penetrated northwards into Transcaucasia from the valleys and low
ridges of northwestern Iran and into Ciscaucasia directly from the Russian
Plain and indirectly from central Asia by way of the northern coastlands
of the Caspian.
The broadest range of the wild ass existed in the Bronze Age. It is
possible that the extinction of the wild ass in Transcaucasia coincides with
the Mongolian invasion in the 13th century, and with the later growth of the
Iranian [Persian] kingdom when the animals of the semidesert were
destroyed in massive hunts. In northeastern Ciscaucasia wild ass survived
until the 19th century.
329
334
ye
ty oe nee 1a Ш
FIGURE 129. Kulans in the Pleistocene semidesert of eastern Transcaucasia
ОЧАГА ГС fT A
Representatives of the families Suidae, Camelidae, Cervidae and
Bovidae inhabited the Caucasian Isthmus in the Quaternary. Pig, deer and
Cavicornia existed there up to the Holocene. Paleontological and
zoogeographical data will be examined in order to analyze the history of
the range development and the evolution of Holocene species of artiodactyls.
Family SUIDAE
The fossil remains of Suidae, abunodontnonruminantfamily, are known
in Eurasia from the Lower Oligocene. The genus Sus, including the extinct
subgenus Microstonyx, dates from the Lower Pliocene. Most of the
fossil remains of Suidae were found in Upper Miocene and Pliocene deposits
of the Mediterranean and southeastern Asia.
Boar —Sus scrofa L. (s. lato). Remains of pigs belonging to the
polymorphic species 5. scrofa occur оп the Caucasian Isthmus from
the Upper Pliocene. The ancestor of the present-day boar was probably
the large Taman boar, 5. tamanensis (Vereshchagin, 1951d).
The next stage of evolution is represented by remains of the Apsheron
boar, S. apscheronicus Burtsch. et Dzhaf., from Middle’ Plerstocene
loams of eastern Transcaucasia, recorded by Burchak-Abramovich and
Dzhafarov (1948).
In the Akhshtyrskaya cave on the Black Sea coast, boar fossils were
found in all strata from Mousterian to late Iron Age, their frequency
increasing with time. Thus, in Mousterian strata they comprised 0.06%,
330
335
in Upper Paleolithic 1.2%, and in роз -Ра1ео с 20.0% of the total
number of osseous remains in each strata.
Boar existed in pre- and post-glacial time in Western Europe (Amon,
1938). North of the Caucasus on the Russian Plain, there have been single
findings of boar in the Upper Pliocene and Middle Pleistocene deposits of
the middle Volga region. During the Pleistocene the boar was probably
very rare even in the river valleys of the Russian Plain, penetrating the
area sporadically from the south in winters with a low snowfall. The rich
fossil collections from the banks of the Don and the Volga contain only one jaw
fragment of a Pleistocene boar (found on the Tunguz Peninsula), while from
the Ural valley only remains of Holocene boars are found (Vereshchagin
and Gromoy, 1952). In general, a number of related facts indicate that
the migration of boars from the Caucasus and their widespread settlement
in the Russian Plain occurred only in the Holocene.
South of the Caucasus, in northern and central Iran, Palestine and Syria,
boar fossils have been found in Paleolithic and post-Paleolithic settlement
strata, although not frequently because of the predominantly dry topography
of these countries.
In examining the contemporary characteristics of central Asian and
Caucasian boar — a concentrated succulent vegetal diet taken from the
soil or subsoil, short legs, relatively poor fur cover and poor adaptation
to low temperatures and deep snow cover — it is evident that the latter
features determined the distribution of this animal in the Quaternary. In
this respect, Formozov's (1946) opinions on the ''chionophobic'' character
of boars is perfectly reasonable. Under the varied climatic and topographic
conditions of a mountain country such as the Caucasus, depth of snow cover,
would not, however, have been such a decisive factor and, in fact, would
have permitted a satisfactory existence for boar on the Caucasus in the
Quaternary.
Settlement strata and burials of the Bronze Age and later cultures on
the Caucasus almost always contain remains of boars. They are especially
common in the early settlements of the swampy Colchis near Anaklia and
in the ancient towns of Kuban. An abundance of boar was characteristic of
the Middle Ages. This is particularly true for the Don valley where
remains of these animals comprise as much as 6% of all fossils collected
in Slav and Khazar strata of Sarkel near Tsimlyanskaya. One can estimate
to a certain degree the distribution and population of boar in the Caucasus
during the historical epoch from art objects and literature. There are no
representations of boars on ornaments and among burial provisions of
the Bronze Age on the Caucasus; these are noted only from the later
Scythian epoch. Spectacular boar representations (Figure 130) are found
in burials of the Kuban area and the Taman Peninsula.
Moisei Khorenskii (1893 edition) mentions the boar hunts of the rulers
of the Arsacid dynasty in the Araks valley in the 1st-2nd century A.D.,
which places them in ancient Roman time. In this description of the reign
of ArtavazdII andthe war with the Romans (Book 2, Chapter 2, page 73)
he said: "Не indulgedinfood and drink: he wandered and rambled on bogs,
in reed thickets, on steep slopes, hunting onagers and boars...'' andfurther
(Book 3, Chapter 55): ''Another time we had a boar hunt among the burned
reeds."
331
336
The decrease in the boar population
and the restriction of its range was
more rapid on the Armenian Highland
because no religious prohibition was
obtained against boar consumption and
because the upland slopes were heavily
deforested. Boar remains which can
now be found on the shallowed shores
of Lake Sevan show how abundant boars
were on this upland in the past. Hunting
was the chief operative factor in the
FIGURE 130. Representation of a boar on a disappearance of boar from [ancient]
bronze plate from the Semibratnoe burial Christian Armenia, a fact that is
(The Hermitage) pointed up by the contemporary
existence of a large boar population
in considerably drier regions of Iran and Turkey. In Georgia the many
mountain forests and almost inaccessible swamps maintained a thriving
boar population over a long time, and it was only the 19th and 20th centuries
that saw the beginning of the rapid extinction of the species.
In Moslem regions, on the other hand, the development of agriculture,
especially rice and melon culture, furnished a nutritional base for a
stabilized boar population. The Lenkoran lowlands and the Samur forests
are examples. In general, the spread of the Islamic population during the
Middle Ages on the Caucasus was, perhaps, one of the primary causes
of the extinction of the range and the population growth during historical
times. Even now, there are more boars in Azerbaidzhan and Dagestan than
in other places. One can judge the abundance of boars on the Caucasus in
the 18th-19th centuries from the many notations contributed by Caucasian
naturalists, students of local lore and travelers.
The rapid decrease in boar population probably commenced in the 1850's
after Russian settlement of the Caucasus. This decline was first observed
on the lower Don, then on the Kuban and later on the Terek and Sulak.
Boars were already rare in the area of the Don estuary and on the Azov
coast by the beginning of the 19th century, and they completely disappeared
about 1812.
In the 1870's boars were still found in the forests near Stavropol (Dinnik,
1914a) from which they have completely disappeared in the 20th century.
Before 1900 the boar population on the northwestern Caucasus declined
considerably because of the growth of settlements of Armenian hunters who
hunted throughout the year.
A small number of boars inhabited the reed thickets of the Kuban as recently
as 1932 in the vicinity of the Cossack villages of Grivenskaya and Petrovskaya.
A local hunter would usually kill no more than ten of these animals per
season.
In the 1930's a small boar population was encountered in the alder-grown
bogs of the lower Rion between Poti and Samtredia — an area which had
swarmed with these animals over many millennia. In general, western
Georgia, Imeretia and Abkhazia supported only a small boar population
by 1930.
In 1909 in the Karayazy forest east of Tbilisi, as many as 50 boars were
killed during one hunting day. In1911, 120boars were killed in a two-day
332
337
period (Kalinovskii, 1900; Markov, 1931а, 1935). Today it is hardly
possible to kill five boars in Karayazy in a day.
In the Kura-Araks Plain the boar population fluctuated throughout the
Quaternary period as the Caspian Sea level rose and fell and created breaks
in the coastal ridges which resulted in the displacement of wormwood
steppes by vast reed—cattail swamps and lakes.
Such a break of the Araks occurred in 1896 when vast floods of Akh-Chala
occurred on the Mugan steppe (Shelkovnikov, 1907). Even today such
adverse factors as the height of the Kura in flood, the extent to which the
steppe lakes fill and the burning-back of reeds largely control fluctuations
in boar population.
The booklet "Agriculture and Forest Economy of the Lenkoran District"!
(1914, pp. 21-22) contains the statement: ''Ten years ago the number of
wild pigs was so high that they became a real menace but when boar hunting
became commercialized, the number of pigs was considerably reduced.
Every year Armenians from the Elizavetpol Province came here to hunt
wild pigs for their skins to use in the manufacture of shoes. Local Russian
hunters kill the swine for their meat. For the last two years, about
5,000 skins were sent to Elizavetpol Province through Lenkoran. "'
During the 1930's the forests of northeastern Karabakh and the northern
slopes of the Lesser Caucasus as far as Georgia did not contain boar.
They were, however, encountered in the Akera-Chai valley and on the
afforested northern slopes of the Zangezur Range where the local population
was replenished by boars migrating through the Araks from the juniper
forests of northern Iran.
Before the later 1940's, a vast tugai forest extended 70 km along the
Kura River from the outlets of the lora and the Alazan to Mingechaur;
it was destroyed preparatory to the flooding of the Mingechaur Reservoir.
The boars which inhabited these tugai forests were either killed or relocated.
In the 1950's, after the closing of the Mingechaur Dam, the tugai forests
below Evlakh began to dry out rapidly and, as a consequence, disappeared
from this area also.
The decline of the boar population in the last decades is visible even in
Talysh where the mountain forests are thought to have abounded in boar.
On the whole, the decline of the boar on the Caucasus during historical
time is a result of its being hunted, both as a valuable trade animal and
as apest. Anthropogenic changes in the landscape and epizootics were of
secondary importance.
The present range of boars on the Caucasus is still important. They are
observed all over the northern slopes of the Main Range and in the foothills,
even near Kislovodsk and Nal'chik. There are few boars in mountainous
Dagestan. The densest population is in the Caucasian Reservation, in the
lower reaches of the Terek and the Sulak, in the Alazan-Agrichai lowlands,
in parts of the Kura and Araks tugai forests and in Talysh (Map 80). In
adjacent territory of Iran, the boar is most numerous on the northern slopes
of the Elburz Range (Sarkisov, 1944e).
The vertical distribution of boar on the Caucasus extends from sea
level to the alpine zone. The piedmont lowlands are inhabited more
continuously; in the mountains distribution depends upon the availability of
seasonal forage and the depth of the snow cover. According to the data of
Donaurov and Teplov (1938), boars are observed in the following zones of
333
338
the Caucasian Reservation (percentage/zone/year): broadleaf forest zone —
48%; dark coniferous forest zone — 39%; subalpine zone — 11%; and alpine
zone — 2%. Е
Table 72 is based upon observations of the author's in 1935-36 in the
Alazan valley and the Zakataly Reservation; percentages reflect fresh traces
of boar (crouching imprints, evidence of digging, and excrements) observed
in each zone or biotope.
TABLE 72, Incidence of boar tracks observed on the southern slope of the Greater Caucasus in Azerbaidzhan
Percentage of total traces observed on prescribed
20-km route
August 1935 October 1935
45 15
32 35
10 32
Zones and biotopes
January 1936
Lowlandsforests of Alazainy: воина eee
Foothillsawith fmitulGes ecnchestifccc Gadd seated & <uces
Oak forest edges of the subalpine zone.........
Alpine MeAadOWS. 7.6 оао suas А eure oe
In Talysh and the Lenkoran lowlands, lacking an alpine zone, the
distribution of boar is somewhat different: in the mountain semidesert zones,
where grain fields and fruit trees are plentiful near the upper forest edge,
the boars concentrate in June and July, while in fall and winter they mainly
inhabit the lowland and the middle forest zone. The daily activity cycle and
distribution of boar are influenced, not only by fodder availability, but by
the degree and kind of hunting. In eastern Ciscaucasia and Transcaucasia,
boars often couch in the vicinity of the railroad (Dinnik, 1914a), in gardens
and among weeds growing in borders of cotton fields (in the Shirvan and
Mugan steppes).
Where they are not hunted or disturbed, boars commence to feed
at dawn and, in the reed thickets of the Kyzyl-Agach Reservation, they
even wander by day. In places where they are constantly harassed, boars
may feed only late at night, and before daybreak will travel sometimes as
much as 10-15 km from their feeding ground to find couching places.
The animal's preferred biotopes in the eastern half of the Caucasian
Isthmus are impassable thickets of tugai, lower forests of Caucasian olive
and willow, bramble shrubs and reed thickets. These thickets are the
primordial biotope suitable to morphological adaptations of the animal.
Any disruptions in the pattern of their horizontal and vertical migrations
in search of food or an escape from deep snows cause the death of
contemporary boars. Dinnik has recorded (1914a, pp. 47-48) episodes of
mass mortality among boars in the 1880's and between 1902-1908 caused by
deep snows, cold and hunger. It seems possible to us that this situation
could have been created by their inability to migrate to the warm Kuban Plain.
They could have reached the Kuban valley from the region of deep snow of
the mountain forests through stands of forest or tugai. But by the 20th
century, their migratory instinct had been suppressed as the result of
334
339
harassment and habitat modifications, such as human settlement and
the deforestation of the piedmont plain.
In the 1940's the total boar yield on the Caucasus was 8,000:
Azerbaidzhan — 4,000; northern Caucasus — 2,000; Dagestan — 1,500;
Georgia — 300; Armenia — 200 (Vereshchagin, 19474).
The future of the boar population of the Caucasus essentially depends
upon the further development of the agricultural and forest economies.
Boars are pests in fields of corn, barley, wheat, rice and melons and
where these crops are grown they will be exterminated or considerably
reduced.
In eastern Transcaucasia the development of irrigation flood-control
and cotton farming will inevitably reduce the boar population through the
reduction of protective cover.
If hunting quotas were established boars could exist in the mountain
forests of the Greater Caucasus and Talysh for an indeterminate time.
Family CERVIDAE
Pliocene strata of the Caucasus have been found to contain remains of
representatives of the genera Eucladocerus, Procapreolus and
Pliocervus, and Cervus, Dama, Megaceros, Alces, and
Capreolus specimens are known from post-Tertiary deposits. Of these,
only Cervus elaphus, Alces and Capreolus have survived to
the present.
Red deer — Cervus elaphus L. (s.lato). Fossils of the red deer
group (C. elaphus) are known in Europe from the Pliocene. From
Mediterranean deposits ten different types of deer are recorded, but only
from the Pleistocene. In the Caucasus the remains of these deer have been
found from the Upper Pleistocene to the present.
For the most part these belongtothe extant C. elaphus maral Ogilby.
The location of the fossil remains (Map 83) indicate the wide distribution
of this species.
North of the Isthmus C. elaphus was as common a settler of the
floodplains and flood valleys in the Pleistocene as the primitive bison and
the horse. In the Novocherkassk Museum its remains rank fourth after
elephant, primitive bison and tur fossils. In riverbed deposits found on
sand banks, bones of C. elaphus account for 3.3% of all the bones
accumulated on the lower Don and 0.5 to 5.0% on the lower Kama and
middle Volga.
South of the Caucasus on the Iranian Plateau remains of Pleistocene
C. elaphus were found in Paleolithic and post-Paleolithic strata of many
caves and in diluvium northeast of Maragheh. It is important to mention
that in caves in the vicinity of Urmia and Behistun bones of deer account
for 57% of the total number of extracted bones. In Mesolithic and Neolithic
strata of the Belt cave in the eastern extremity of the Elburz Range there
were few deer remains — only 0.8% (Coon, 1951). In the Neolithic strata
of the ancient townof Annauin western Turkmenia deer remains are rare
(Duerst, 1908). C. elaphus was a common food of Paleolithic hunters
in Lebanon, Syria and Palestine. In spite of its frequent occurrence,
there can be no doubt that the habitation of deer in southwest Asia was
confined to river valleys and wooded slopes from at least the Pliocene.
335
340
341
Fossils of С. elaphus are аз numerous in postglacial deposits as
they are in Pleistocene deposits. In the Holocene loams of the Stavropol
and Pyatogor'e areas, inthe Yegorlyk, Kumanand Podkumok river valleys, and
in talus fragments of the slopes and banks of Lake Sevan on the Armenian
Highland, skulls, teeth and antlers are especially common.
These findings (see Chapter II) indicate that deer was widely distributed
over the Ciscaucasian steppes and Transcaucasian plateaus, at least in
the spring and fall and at the beginning of winter.
This earlier distribution of deer on the steppes should not be associated
with a wide deforestation of these areas as Shelkovnikov (1930), Dal' (1947a)
and others thought. In the spring—summer season deer do not require large
forests; on the contrary, they migrate from the forests to steppe and
meadow areas to escape from bloodsucking insects.
At present deer inhabit the Don, Volga and Ural valleys and adjacent
steppe and forest areas. Doubtless the Caucasian population of C.elaphus
was always related to the Eastern European population. In the strata of
the Middle Ages in the Sarkel fortress near Tsimlyanskaya C. elaphus
bones comprised 4.3% of all the fossils investigated.
During the post-Paleolithic C.elaphus was the favorite subject of
painters and sculptors of the Caucasus; it was used in the decoration of
pottery, on arms and on clothes. Beginning with drawings on rocks
in Kabristan (Figure 131), the deer subject is repeated in bronze pendants
of Kobanian culture, in the ornamentation of Khodzhala and Sevan bronze
belt buckles and especially in gold articles of Scythian-Sarmatian culture
of the Trans-Kuban Plain and south Ukraine (Figures 132-133). Deer figured
importantly in religious belief and rituals. According to legends of the
Abkhazians and Imeretians, deer coming on the Feast of St. George to
caves or chapels dedicated to the saint were killed for feasting (Dzhanashiya,
1915). In a number of Ossetian dzuars antlers and sacrificial skulls of deer
predominate among the remains of other wild, hoofed mammals. All this
indicates that the deer population during the postglacial period did not
decrease as rapidly and catastrophically as did that of tur and bison.
Osseous remains of deer found in Quaternary deposits of steppe regions
are often cited by naturalists as evidence of either a former steppe way
of life or an earlier existence of forests in steppe regions. Recent
observations of deer behavior and feeding habits and of the structure of
the lower molars clearly indicate that the C.elaphus of Pleistocene and
historical time was an inhabitant of the steppes, especially in spring and
fall. During these seasons they grazed on the young grasses of the steppes,
and at other times of the year inhabited the floodplains and oak forests
in ravines. Deer were widely distributed in steppe and forest-steppe at
least until the late Middle Ages. Guillaume le Vasseur de Beauplan
(1832 edition, p.92) noted flocks of deer in the Ukrainian steppes at the
beginning of the 17th century: ''Deer, fallow deer and saiga are
encountered in flocks in the Ukrainian steppes.'' The present existence of
deer on the steppe-pastures of the Askaniya-Nova Reservation gives
credibility to this earlier report.
Shal'kovskii (1885, p.189) also noted in his account of 18th-century
Zaporozh'e that hunters ''also caught hares, wolves and deer which, until
1760, inhabited the marvellous forests of the Trans-Dnieper Zaporozh'e. "'
336
(340)
FIGURE 131. Beyuk-Dag rock south of Baku showing representation of a deer hunt
Photograph by the author, 1946
FIGURE 132. Kobanian artifacts from North Ossetia (Uvarova, 1900)
1 — bronze rod representing a deer hunt; 2 — deer head
337
342
During historical time а marked decline in the deer population can be
traced to the development of cattle herding and agriculture. The greatest
decline in the deer population of the East European and Caucasian steppe
and forest-steppe zones, however, probably occurred when horses were
domesticated and used in stag hunting. During the campaigns of the
Scythians, Huns, Kipchaks and later the Mongolians, deer living on the
steppes were rapidly decimated. It is possible that stag hunts forced the
retreat of deer into the forests and forested foothills where they were out
of reach of the mounted hunters. At the beginning of the 15th century,
Josaf Barbaro (1836 edition) witnessed a Tartar ''pagany'' — mounted
hunt — for bustards and other animals which formed an encirclement of
120 miles (!) "апа [зам ] 4еег and other animals escaping from the Tartars.'"'
The first phase in the depletion of C. elaphus during historical times
was the extermination of its steppe population.
The second phase of decline began in the late Middle Ages as a result
of agricultural development and the rapid eradication of forests. This forest
extermination was accompanied by desiccation and subsequent dwindling
of land in the Caucasian foothills and high plateaus.
Historical data on the distribution and yield of deer have been preserved
since the 16th century. Georgian and Iranian chronicles and the ancient
literature of Georgia, Azerbaidzhan and Iran mention the deer hunts.
Thousands of peasant beaters and warriors took part in hunts organized
by the Persian Shahs during the late Middle Ages in the forests of Gilan
and Karabakh.
The populousness of deer in Mazandaran and Gilan in the 16th century
can be reckoned from an end-of-the century account by Adam Olearius
(1870 edition, p.730): the Shah Takhmasp and his retinue killed 2,000 deer
and goitered gazelles — ''agu'' — and he ordered the "building of a tower
from their antlers."
FIGURE 133, Scythian artifact from the Trans-Kuban Plain
Gold-plated reclining deer from a hill near the Cossack village of
Kostromskaya
338
343
In "Тре Geography of Georgia" (1904), Prince Vakhusti reports that,
at the beginning of the 18th century, the 94th czar, Vakhtang VI, and his
entourage killed 180 deer in one day on the Trialet ridge southwest of Tiflis.
The historian Sekhnia Chkhaidze writes in ''The History of Georgia"’
(1945, p.408): ''The Czar Vakhtang Levanovich (1703-1721) invited persons
of high rank to a hunt in Trialetia and Shanbiani. The first day 60 deer and
countless other animals were killed."
At this time deer still inhabited the reed-grown and forested floodplains
of the Don, Kuban, Manych, Kuma, Kura, Terek and Sulak rivers.
The disappearance of deer from Ciscaucasia in the 17th and 18th
centuries can be attributed to the settlement of the country by Cossacks
and the widespread use of firearms. Dinnik (1914a, p.97) reports that he
found only fragments of antlers as evidence that deer existed there
"60-70 years ago,''i.e., in the 1850's and 1860's.
In the environs of Krasnodar and on the right bank of the Kuban, where
they had been protected, deer still lived up to the early years of this
century, when they were finally exterminated during the Civil War.
According to Dinnik (1914a) the northern limit of their range on the
northwest Caucasus passed through the Cossack villages of Peredovaya,
Kaladzhinskaya, Makhashevskaya and north of Maikop. The largest deer
population was observed in the upper reaches of the Belaya, Bolshaya and
Malaya Laba rivers where 20-30 deer could be encountered in one day. In
winter, deer ranged as far as Novorossiisk and Kabardinka. Later, the
distribution area shifted southward. In the early 1930's, the northern range
limit lay in the foothills along the line of the villages of Solenyi, Buguzh,
Dakhovsk, Samursk, Neftyanaya and Navaginskaya. The range reached its
western limits in the upper reaches of the Pshish River (Nasimovich, 1936a).
In the 1940's deer could still be seen east of the Caucasian Reservation
near the railroad stations of Predgradnaya and Mikoyan-Sakhar.
Rossikov (1887) tells of hunting deer on the piedmont plain north of
Nal'chik as late as the 1870's.
FIGURE 134. Illustration of deer hunt in Digoria from "polati" [sleeping ledges or bunks] in the
Digorized cave
Photograph by author, 1947
339
344
The last of the deer in the piedmonts of North Ossetia were exterminated
(Figure 134) by the 1920's (Vereshchagin and Naniev, 1949). They survived
for a longer time on the Sunzha tributaries near Gudermes and they are
still found in tugai forests along the Sulak and Terek on the Dagestan plain.
In the Dagestan uplands deer were numerous in the gorges of the Avar
Koisu. Many antlers have been found in burials in deciduous and coniferous
forests near Antsukh [Bogaz Range] and Tlyadal, and in Khevrusetia [eastern
Georgia] and Tushetia [Andi Koisu region] (Dinnik, 1914a; Maruashvili,
1955). The deer migrated to Dagestan from the southern slopes of the
Greater Caucasus only in the summer to escape from bloodsucking flies,
returning again in October or November to winter on the southern slopes.
It is the practice of the Dagestanians to hunt them in the passes from
Dagestan.
Deer had disappeared from the Samur-Khachmas lowland forests by the
beginning of this century and, according to old-time residents of the
village of Kusary, they were gone from the higher forests of the Kuban area
by the 1880's.
In the Kura lowlands they could still be encountered in the tugai forests
and reeds of the middle and lower reaches of the Kura River as late as
the end of the last century. Older residents of the ancient village of Shil'yan
and of the Shirvan steppe recall deer living along the reed-grown margins
of the Lake Shil'yan, and a path through the reeds is still called ''mara]l
се!" or ''maral elu'' (deer path). Deer was already rare in Talysh by the
time of Radde, Satunin (1905b) and Dinnik (1914a). According to the
Azerbaidzhan Game Inspector, V.V. Vitovich, the last deer in Talysh was
killed in 1912.
In Karabakh, deer had disappeared from the Dzegam and Shamkhor
ravines by the 1920's. Melik-Shakhnazarov (1898) reported that in the 1890's
in Zangezur deer were so abundant that some hunters killed 150-200 head
a year. Osipov (1898) tells of deer being common in his time in the vicinity
of Chaikent (south of contemporary Kirovabad).
[п {Бе 1930's and 1940's some deer still survived in the Karayazy forest
on the Kura River where in the 1890's 15-20 specimens had been killed in
one day.
In western Georgia, deer were either exterminated or driven into the
mountains from the lower Rion, Kodor and Ingur by deforestation at the
start of World War I.
The present range of C. elaphus is comprised of four or five isolated
tracts (Map 83), the largest of which is located in the northwest and includes
the Caucasian and Teberdinskii reservations in the Pshish, Kurdzhips,
Belaya, Laba, Malyi Zelenchuk, Bolshoi Zelenchuk and Kuban river basins.
In the summer, most of the deer inhabit open places in the fir forests in
the interior of the Caucasian Reservation at altitudes of 1,700-2,200 m.
Their winter habitats are found in sunny places on the slopes and in the
valleys of the lower fir belt and the beech-forest zone with a snow cover
of 0.3-0.5 т. After the sharp decrease during the Civil War, the deer
population commenced t9 rise again. A 1934 census (Nasimovich, 1936a)
showed a total stock of 900 head. By the 1940's, there were at least
3,000 head of deer in the Caucasian Reservation.
In northeastern Ciscaucasia deer survived in forests and reed- grown
areas of the lower Terek from the Cossack village of Chervlennaya to
Kizlyar, and on the Sulak from Khasavyurt to the lower river.
340
345
These deer live the year around in the tugai stands of the lowlands,
because there are no longer any alternative locations open to them for
migration: the northern foothills of Dagestan are now deforested and
populated.
In western Transcaucasia a small deer population still inhabits the
forests on the middle course of the Ingur. In the beech forests of the
Borzhomi Reservation (on the eastern spurs of the Adzhar-Imeretia
Range), El-Mar (Е. 1.. Markov, 1939) counted almost 900 head of deer in
1934. The summer habitat of the Borzhomi deer had previously been
located on the high, cold, deforested plateaus of the Dzhavakhetia and
Akhalkalakhiuplands. In eastern Transcaucasia deer are found in the
Karayazy forest and on the southern slope of the Main Range from the upper
Alazan to the Gerdyman-Chai. The largest deer population has been
observed in the Zakataly Reservation in the Belanukh-Chai and Katekh-Chai
basins; it was estimated in the 1940's at 1,000 head. To the east, in the
Kutkashen and Ismailly districts, deer are numerous both in the mountains
and on the plain, and can be encountered near Chukhur-Kabala, Ismailly,
Kalandzhak, Rushan-Kend and other villages.
Our own observations made in the 1930's showed the summer habitat
of deer to be on the southern slopes of ridges in the upper third of the
beech forest zone and in the subalpine zone. If there were no cattle or
dogs, they would venture onto the pastures. In the winter they inhabited
beech forests in the middle altitudes of ridges with southern exposures.
During winters with a heavy fall of snow deer sometimes appear on the
open hilly steppe south of the Shemakha. This habit and the year-round
existence of the Sulak population in reeds and tugai explain why deer remains
were found together with those of ass and saiga in Pleistocene deposits
on the Apsheron Peninsula. Notwithstanding its earlier history and these
present variations, the contemporary C. elaphus must be regarded as
а mountain-forest species, not accustomed to remote migrations to the
plain. The differences between various deer populations in habits and in
the directions their migrations take merely testify to the great adaptability
of the animal and require a correct reading of its contemporary behavior
as a response of its higher nervous system to human influence.
During the 1940's poachers on the Caucasus averaged an annual bag of
200-250 deer (Vereshchagin, 1947а).
The Caucasus is the chief deer preserve in the U.S.S.R. The subsequent
fate of the species in this region will depend upon the effectiveness of
measures taken for their protection. If poaching is controlled, it will
allow renewed breeding in the forests of Karabakh, Talysh and the Kusary
sloping plain. In addition, it should be possible to resettle Caucasian deer
in the future in the floodplain and watershed forests of the Russian Plain.
Fallow deer —Dama cf. mesopotamica Brooke. Fossil remains
of fallow deer similar to the Mediterranean D. dama L. are recorded
from Pliocene and Pleistocene deposits in Italy, England and Germany.
A fragment of mandible and a fragment of antler discovered on the
Caucasus in Pleistocene alluvium at Adzhi-EHilas near Yerevan are unique
and can be reported here as D. mesopotamica only conditionally.
In southern Asia Minor and Palestine fallow deer was the most frequent
bag of Paleolithic hunters (Bate, 1937).
341
Artistic representations of Mediterranean
fallow deer heads in bronze (Figure 135) were
found in the Semibratnoe Scythian burials on the
Taman Peninsula, and also in burials in the
southern Ukraine. This is a basis for presuming
that the distribution areas of the two species,
i.e., Mediterranean and Mesopotamian,
encompassed the southern Balkan Peninsula,
the Black Sea area, Asia Minor, Irag and Iran
(Map 82). The fallow deer is an inhabitant of
broadleaf forests and thickets and of the
undergrowth of Mediterranean river valleys.
The species is more thermophilic than Cervus
elaphus.
By the beginning of the 20th century, the
pega age ооо, fallow deer was nearly extinct in its natural
aide saree na Ae range; it has since been bred under protected
Scythian burial (The Hermitage ) conditions throughout Western Europe and north
Africa (Ellermann and Morrison-Scott, 1951).
A successful experiment in breeding fallow
deer was carried out on the Caucasus near Borzhomi before the revolution
(see Chapter VI).
Through future regulation of hunting, the fallow deer can become a
worthwhile element inthe piedmont shrub-forest zone of the Caucasus.
It can populate the forests of Stavropol, the Dagestan piedmont, eastern
Armenia, the Nakhichevan A.S.S.R. and the Nagorno-Karabakh uplands.
346
FIGURE 136. Golden plates from Scythian burial with representations similar to giant deer.
Berlin Museum (Bachofen-Echt, 1937)
Giant deer —Megaceros euryceros Aldr. Upper Pliocene remains
of this deer have been found throughout Eurasia.
In the Pleistocene the distribution area of M. euryceros included
the forest-steppe and the steppe zones of Europe and Asia. Its remains
342
347
are езрес1аПу common in Khazar strata of the Volga valley, where they are
found on the banks and comprise 2—8 % of all the bone material. They are
also common in Pleistocene strata of Western Europe. On the British
Isles fossils have been found in caves, diluvium and peat beds. Some
paleontologists, reasoning from 12th-century drawings and the rather fresh
remains found in Irish peat beds, view М. euryceros as an inhabitant
of Ireland up to the time of the present era (Mitchell and Parkes, 1949). The
European continent lacks such recent remains indicating that the species
was probably extinct here in the Upper Pleistocene. However, Pidoplichko's
report (1951) of a find of M. euryceros remains inthe U.S.S.R., which,
according to the bone calcination, showed a Recent origin, deserves further
attention and verification.
Kitchen middens containing M. euryceros remains, whichare common
in Paleolithic caves of the Crimea, are completely unknown from the caves
of Palestine, Syria and Lebanon. On the Caucasus Isthmus, fossils have
been found from the Upper Pliocene — on the Taman Peninsula, through
the Mousterian — in Abkhazian caves and camp sites on the Trans- Kuban
Plain (Map 82). M. euryceros inhabited Abkhazia in a mountain
mesophytic environment.
The species is rarely found in Pleistocene strata of the eastern
Transcaucasian steppes, andin the Binagady asphalts only one bone among
tens of thousands was identified as M. euryceros. No remains of giant
deer have been found in Holocene deposits, even those as large as the Lake
Sevan shore and the Sarkel strata of the Middle Ages. It is not depicted in
any Caucasian Paleolithic or post-Paleolithic drawings and scuptures.
Stylized representations of Cervus elaphus in gold plate taken from
Scythian burials on the Trans-Kuban, which somewhat resemble representa-
tionsof М. euryceros (Figures 136, 137), led Bachofen-Echt (1937) to
the erroneous assumption that giant deer existed on the Caucasus in the
first millennium B.C.
FIGURE 137. Skull of Megaceros euryceros from the environs of Rostov-on-Don
The usual osseous deposits containing M. euryceros found in flood-
plains and peat beds reveal an ecological similarity to the elk. The
structure of metapodia and especially of hoof phalanges place the species
in a position intermediate between elk and ox. It could probably endure
343
a deep snow cover, but could not undertake long marches in the snow like
the elk. It was an inhabitant of the cold forest-steppe and the river valleys
of the plains, living in floodplain forests and thickets together with
Elasmotherium,and Elaphus irogontherii,
Caucasian elk — А1сез alces caucasicus М. Ver. Remains of
elk of the A. alces L. group are recorded from Pleistocene deposits of
the northern Holarctic, but are rarely reported south of the 50th parallel.
In the western Mediterranean Pleistocene deposits seldom contain
elk, although other species of the Alces genus are sometimes so identified
(Wolf, 1938; Pidoplichko, 1951 — map, p.79, reports all findings as
A. alces). Earlier reports on finds of elk remains in the Paleolithic of
Palestine, Syria and Lebanon (Blanckenhorn, 1910) were in error (Picard,
1937). The same may be said more emphatically for Egypt. North of the
Caucasus remains of Pleistocene elk have been found at Borshevo II, the
encampment site near Voronezh, and on scarps of the Volga bank near the
village of Osypnoi Bugor beyond Astrakhan. The species does not occur in the
Crimean Paleolithic. On the banks of the middle Volga and especially of
the lower Kuma, elk remains generally account for 5% of all the large-
species bones.
On the Caucasian Isthmus elk remains are recorded for the Upper
Pleistocene (Upper Paleolithic) from cave strata in western Transcaucasia
from Khosta to Kutaisi.
The Mousterian strata of the Akhshtyrskaya cave, the Il'skaya
encampment and the Apsheron Peninsula asphalts revealed no remains
of elk. It would be justifiable to conclude that the elk first penetrated the
Caucasus from the north in the Upper Pleistocene, although it is equally
possible that the development of the ancestral form could have taken place
on the Caucasus, or, for that matter, anywhere in the eastern
Mediterranean. In either case, it is a matter of established fact that the
elk survived the Upper Pleistocene on the Caucasus and became extinct
only at the beginning of the 19th century.
The presence of elk on the Caucasus during the Bronze Age can be
assumed from bronze buckles and pendants of Kobanian workmanship found
in archaeological excavations in North Ossetia (Uvarova, 1900, p. 80,
348 Figure 75; Vereshchagin, 1949a). In the collections of the Moscow State
Historical Museum there is a bronze buckle with a design of two elk heads
which was excavated near the village of Atage in the Sunzha valley and
belongs to the second period of Koban culture (Figure 138). An interesting
relief resembling an elk is carved ona stone from Euyuk, a Hittite town
which existed in the first half of the second millennium B.C. in northeastern
Asia Minor. This suggests the possibility that south of the Caucasus elk
inhabited forested tracts in Asia Minor in historical time.
Representations of elk appear in gold and bronze (Figures 138, 139) and
on the ornamentation of quivers found in burials on the Trans-Kuban
piedmont plain and in the southern Ukraine.
Finally, Keller (1909) reported the find of gold pendants depicting elk
in ancient Greek settlements on the northern shore of the Black Sea.
All the elk representations of the Scythian epoch are of great importance,
not only in reconstructing the early range of the elk, but alsoin assigning the
origin of gold and bronze artifacts from Scythian burials with ornamental
and relief designs of animal subjects. Taking into account that the southern
344
margin of the elk range in the Holocene passed through the Caucasus and
the northern Black Sea area, it becomes apparent that most of these
Scythian artifacts were in fact produced at their cultural source — in
Scythia — rather than, as often assumed by historians and archaeologists,
in the southern Balkans, Asia Minor or the Aegean Islands.
t , , Л yom
FIGURE 138. Bronze buckle with
elk head design from the village of
Atage on the Sunzha (Moscow State
Historical Museum )
FIGURE 139. Bronze plate
depicting an elk's head (1:2)
from the southern Ukraine
There are references to Caucasian and
southern European elk in the literature.
Jean de Luc (1879 edition) in his
description of a journey -nto the Nogai Tartar
territory at the beginning of tne 17th century
tells of seeing elks, in addition to wild horses,
deer and other animals. Clarke's reference
(1810, p. 386) to two young and very tame
"elks'' which he saw near Ek.terinodar should
certainly be assigned to deer rather than to
elk, and Kirikov was wrong to use this citation
(1952) as proof of the earlier existence of elk
on the Caucasus.
Pallas (1831)reportedin his 'Zoography"
that elk was found on the Caucasus, and
Lul'e (1875), in his description of the beliefs
and customs of the Circassians, wrote of
Mezitkh, their god of the forests, who rode
on a boar with golden bristles. At his nod,
"deer and elks assembled in the forests, and
then young maidens milked the does. "'
The presence of elk in the contemporary
fauna of the Caucasus was later denied,
however, by the best zoologists (Dinnik, 1896,
1914a; Satunin, 1903b). Satunin wrote:
'Thus he (Pallas) placed even the elk (Alces
а1сез L.) оп the Caucasus, although there
can be no doubt that this species never
inhabited the Caucasus."' In our time, Kulagin
(1932) and Buturlin (1934) expressed doubts
about the occurrence of elk on the Caucasus.
Kulagin's reservations, which he bases on
Polferov's story, ''Eznos'', are really only
tenuously connected with it. In fact, Polferov
mentions encounters with elk and capercaillie
somewhere in the taiga zone, and not on the
Turkish border as Kulagin surmised.
In the 1930's elk bones were found in
archaeological excavations of 8th—12th-century
towns near the Cossack village of Tsimlyanskaya on the lower Don.
Figure 140 shows a semi-fossilized skull of ancient elk identified by the
author in the Stavropol Museum collections in 1944. While investigating
Ossetian dzuars in the Urukh Basin in 1947, we found seven elk skulls
(Figure 141). These findings establish that the elk inhabited the foothills
of the northern Caucasus contemporaneously with the primitive bison,
both becoming extinct at the beginning of the 18th century (Vereshchagin
345
and Semenov-Tyan-Shanskii, 1948; Vereshchagin, 1949а, 19495, 1955;
Vereshchagin and Ganiev, 1949).
FIGURE 140. Semi-fossilized skull of elk in the Stavropol Museum
Photograph by author, 1945
Ritual collections containing remains of elks, similar to those of Digoria,
are probably to be found in other gorges of the northern slopes of the
Greater Caucasus, particularly on the Balkar-Cherek.
350 Even during the time when the Ossetian dzuars of Digoria flourished and
skulls and horns were being accumulated in them in quantity, elk was
already rare by comparison with deer and even bison. The seven skulls of
elk found in our investigation of 16 dzuars were in a 10% ratio to bison
skulls and 0.8% to deer skulls. The extinction of the species was brought
about through direct extermination and through the general destruction of
forests on the Terek-Sunzha plain. It is possible that the last of the elks
survived until the beginning of the 19th century.
During the 1950-1951 excavation of the Sarkel fortress on the lower Don
carried out by The Hermitage (Artamonov, 1952), kitchen middens were
found to contain manyremains ofelk, together with remains of deer, boar,
saiga, bear and beaver. The elk remains comprised 0.3-0.8% of all the
bones of domesticated and wild animals found during the excavations. *
Elk remains were also foundrecently, together with those of reindeer, inthe
Roman strata of Olvia (Pidoplichko and Topachevskii, 1953).
A.A. Sadovskii has reported (through a verbal communication) his
observation in 1937 of a mandible of elk among a large quantity of wild
ungulate bones, including deer and roe deer, which had been uncovered
during the excavation of the ancient burial of Naokhvami in Mingrelia.
The relict alder swamps in the lower reaches of the Rion with thickets
of osier and buckbean are even today an ideal elk habitat.
* К.В. Yur'ev, G.V. Khrabrov and others investigated 200,000 bone fragments under my direction in Sarkel
in 1952-1953.
346
351
пы весе 5 1емое voile Ш 1 ВЕ
Trans-Kuban was confirmed by the find
of a horn (Figure 142) in the pebbles of
the Urup River near the Cossack village
of Otradnaya. The horn is preserved in
the Krasnodar Museum; although wind-
eroded on the surface, the inside shows
it to be quite recent.
If the elk settled on the Caucasus from
the north during the Upper Pleistocene,
it can be assumed that the principal
penetration route of the species passed
southward through the valleys of the Don
and the Volga. However, the possibility
of an immigration route from the southwest
through the Balkans and Asia Minor cannot
be excluded. The distribution of elk on
the Caucasus was probably limited by the
foothills (Map 84). Their chosen biotopes
(Figure 143) were along spring and swamp
rivers of the Trans-Kuban and Terek-Sunzha
plains, on the littoral terraces of Abkhazia
and in the lowlands of Colchis. As
opposed to С. elaphus, the elk did not
adapt to the Caucasian Range and to the
mountain-forest environment.
When the lumber and hunting industries
of the country are regulated, it will be
possible to resettle elk on the Trans-Kuban
sloping plain in the valleys of the Afips,
Psekups, Laba and Zelenchuk rivers.
It is also possible that the species might
be successfully settled in young mountain
forests of pine, aspen and willow in the
Teberda and Caucasian reservations.
FIGURE 141. Skull of elk from the Roe deer —Capreolus capreolus
Digorized cave ie wand Crcapreolus pyoargus Pall.
The genus Capreolus is of Middle
Pliocene origin (Simpson, 1945). The
ancestor of the contemporary genus is
probably Procapreolus which is known from the Lower Pliocene of
Europe and Asia. Flerov (1952, pp. 98-99) considers that the contemporary
roe deer is the descendent of Mio-Pliocene Cervulinae.
The infrequently found remains of small European roe deer derive from
various types of Quaternary deposits in Europe and Asia, almost all of
them within the limits of the contemporary range (Wolf, 1938; Pidoplichko,
1951). For the Caucasus, roe deer remains are recorded from Upper
Quaternary travertines of Mount Mashuk and from Transcaucasian cave
strata, Acheulean to Recent, at Kudaro II, Akhshtyrskaya and Gvardzhilas
caves and others. A large form of С. pygargus has been recorded from
further south for Palestine, Syria and Lebanon from Acheulean to Neolithic
cave strata (Picard, 1937; Bate, 1937).
Photograph by author, 1948
347
FIGURE 142. Horns of elks:
1 — from the strata of the Sarkel fortress on the Don; 2 — from the pebbles of the Urup River
On the Russian Plain remains of a large roe deer were found in Upper
352 Paleolithic strata of Kostenki on the Don and in Upper Quaternary alluvium
of the middle Volga region (Khryashchevka). Tothe west, inthe Ukraine and
the Crimea, only remains of smaller specimens were discovered in
Paleolithic strata. Roe deer remains are numerous in Holocene deposits
of the Caucasus, e.g., in Copper—Bronze-Age strata of camp sites near
Anaklia, in post-Paleolithic strata of the Akhshtyrskaya cave, etc. (Map 85).
During the excavations of the Sarkel fortress on the Don remains of the
large Siberian roe deer were found in 8th—13th-century strata. The North
Ossetian dzuars, e.g., the Digorized cave, contain preserved skulls and
horns of small roe deer of C. capreolus type hunted in Urukh between
the 15th and 20th centuries.
Roe deer is seldom depicted on artifacts left by the ancient Caucasian
tribes. One of the rare examples is a silver bucket from a Bronze Age
burial onthe Trialet ridge which bears a contour representation of a roe
deer wounded in the chest by arrows (Kuftin, 1941).
The range of roe deer on the Caucasus in the Recent has continued to be
widespread, and today the species is better preserved than other ungulates.
A larger form, similartothe Siberian, inhabits the northern Caucasus
(Dinnik, 1914a), and a small form is found in Transcaucasia. The roe deer
does not occur in the reedlands of the lower Don and Kuban. It is
1704 348
certain that, as late аз the Middle Ages, the Caucasian range of the large
roe deer was connected with the range of the large Siberian roe deer along
the Don valley. The presence of the large roe deer in Ciscaucasia can
probably be explained by a relatively recent postglacial migration from
the north.
FIGURE 143. Caucasian elk
The ultimate disappearance of roe deer from the island forests near
Stavropol took place in the 1920's. There are few of these animals
remaining in the broadleaf forests of the Trans-Kuban sloping plain between
353 Khadyzhensk and Maikop. But on the northwestern spurs of the Greater
Caucasus, they are still populous in beech—hornbeam forests (e.g., near
the Goryachi Klyuch and Krymskaya).
TABLE 73. Distribution of roe deer in the Zakataly Reservation based on number of tracks encountered
daily in summer and winter
Southern spurs; beech—
hornbeam and aspen
forests with glades;
700-1,800 m
Upper forest belt and
subalpine meadows;
2,000-2,400 m
Deep gullies in
ravines; beech forests;
1,200-1,600 m
Date
22-26 August 1935
21- 26 November 1935 (snow:
15-30 cm)
1 January 1936 (snow:
30-50 cm and more) 5
349
354
In Kabardino-Balkaria and North Ossetia, roe deer inhabit the forests
of the Chernye Gory, the canyons of the Skalistyi ridge and the sparse
forests on the northern slopes of the Terek Range. They are not found in
the Sunzha valley, but have been observed in the tugai along the Terek and
Sulak between the Cossack villages of Chervlenaya and Kizlyar. The forests
of the piedmont spurs of the Grozny Region contain a roe deer population,
but the area near Makhachkala does not, and the many wolves in the
reedlands of the lower Sulak and Terek preclude the habitation of roe deer
there. Neither are they numerous in the forests of the Kusary sloping plain.
On the whole, the distribution of roe deer on the northern slopes of the
Caucasus reaches its summer limits at altitudes of 1,500-2,000 м. In the
winters when the snowfall reaches disastrous proportions the animals
descend to the piedmont plains, sometimes even venturing into the villages
of the Maikop area (Olenich-Gnenenko, 1955).
In western Transcaucasia, the population of roe deer is sparse in the
lower forest belt because of the dense network of villages; it is heavier in
the middle third of the forest zone. The snowfall in the upper Svanetian
gorges, e.g., onthe Ingur, is too heavy to support the existence of roe
deer there:
A small population inhabits the Colchis alder swamps near the Chaladidi
station and on the middle course of the Pichora River, where Poti hunters
kill as many as 50 animals yearly. Roe deer is common on the northern
slopes of the Adzhar-Guri Range and the Trialet ridge. Оп the southwest,
its range extends to Asia Minor. In Palestine roe deer was still common
in the last century in the Carmel Mountains where the last specimen was
killed in 1912 (Bodenheimer, 1935).
In east Transcaucasia it is found on the southern slopes of the Main
Range inhabiting mainly the lower third of the forest belt where there are
thickets of Cornelian cherry, hawthorn and hornbeam, or the glades of
beech—hornbeam forests overgrown with giant fern and blackberry.
In 1935 we made the following observations of fresh roe deer tracks on
rivulet shoals and in the snow in the Zakataly Reservation as they were
distributed altitudinally along the Tala-Chai and the Katekh-Chai. (Table 73).
On the Alazan-Avtaran lowlands the habitat of roe deer is confined to
the oak—C aucasian-wing-nut forests, and the population is greater in winter
than in summer. The species doesnotinhabit the tugai, reedlands and
oases of the Kura-Araks lowlands.
On the Lesser Caucasus roe deer can be found wherever there are large
stands of deciduous forest. In summer it inhabits subalpine tall-grass
meadows near relict forest islands at altitudes of 1,800 and 2,000 m in the
gorges of the upper Debed and Akstafa rivers.
In Karabakh many roe deer are to be found in the Terter-Chai beech
forests. They are not encountered on the southern slopes of the Armenian
Highland.
In Talysh and in the forests of the El'brus Range, the few deer live in
isolated colonies in well-defined areas of some of the gorges. Only single
tracks could be observed in the Vassaru-Chai and Vilyash-Chai ravines in
our investigations during June and July of 1945.
Broadly stated, roe deer is frequently found on the Caucasus, but never
in the numbers in which it is found in the piedmonts of northeastern Tien-
Shan or the Maritime Territory.
350
355
In the Iranian uplands, roe deer are sparsely settled in relict forest
islands. According to Blanford (1876) roe deer inhabited only the forests
of Asterabad. The southern margin of the species range probably
lies in the passes of northern Mesopotamia.
Although the principal range of roe deer is now located in northern
Eurasia, it can certainly be regarded as an Upper Pliocene species of the
Caucasus whose closest ancestors are to be found in the Lower Pliocene
of Transcaucasia and throughout the Mediterranean.
The roe deer is adapted to a forest habitat and a diet comprised mainly
of leaves; it has the capacity for very swift movement and high jumps in
shrubs and tall grass; it does not endure deep snow.
The rapid decline of the roe deer population and range on the Caucasus
occurred in the last century as a result of unrestricted hunting and
deforestation (see Chapter VI).
The present yield of roe deer is estimated as follows: northern Caucasus
—- 4,600; Azerbaidzhan — 1,500; Georgia —=|700; Armenia — 300; Dagestan —
200 (Vereshchagin, 1947d).
The development of agricultural shelterbelts in the Ciscaucasian plain
will undoubtedly create ample opportunities for the breeding of roe deer
in a cultivated landscape.
Family BOVIDAE
The Quaternary fauna of the Caucasian Isthmus includes representatives
of the sub-families Bovinae, Antilopinae and Caprinae. Seven genera occur
im@rhnes Holocene faunareGaziell agg Siaiean) (Ru piiciarp nay Сара
@vas) eos, and) Bilsion,
Goitered gazelle —Gazella subgutturosa Со. Fossil remains
of anumber of Gazella species are known from the Upper Miocene- Lower
Pliocene. The genus is generally regarded as a Miocene species. Fragments
of horn stems and teeth of gazelles, which show some similarity
to the Mediterranean Pliocene G. deperdita Сегу., have been found on
the Caucasus in Sarmatian deposits near Georgievsk, in Middle Pliocene
sandstones south of Baku and in Lower Quaternary strata of the Taman
Peninsula.
In addition, a fragment of horn core which resembles goitered gazelle
horn was found in postglacial loams in the upper reaches of the Kalaus,
southwest of Stavropol (Ryabinin, 1918).
Remains which are true to goitered gazelle type were reported for
Pleistocene strata of the Tamtama and Bisotun caves in central Iran (Coon's
excavations, 1951), for Mesolithic strata of caves near Dzhebel and
Krasnovodsk (Okladnikov's excavations, 1949) and for Anau strata near
Ashkhabad (Duerst, 1908).
Picard (1937) and Bate (1937) have recorded remains of gazelle similar
toy G. pame ldagPall исчо саит ааа Gh isu bg utitium@siayircom
Paleolithic cave strata (Acheulean to Upper Neolithic) of Lebanon, Syria
and Palestine. No fossil remains of goitered gazelle have been found in
Pleistocene deposits of the Russian Plain nor in the Middle and Upper
Pleistocene of eastern Transcaucasia. Validated finds of goitered gazelle
remains on the Caucasus are recorded only from Holocene strata of the
historical age in Ciscaucasia and Transcaucasia (Map 87).
351
356
Goitered gazelle remains are particularly numerous in the strata of
ancient Baku in which they account for 14.5% of all the bones found. Their
ratio to sheep and goat bones is 3.9:10.
Most of the present range of goitered gazelle lies within northwestern
Iran and Afghanistan and the deserts of central Asia.
The absence of goitered gazelle in the Binagady fauna of the Middle
Pleistocene and the frequency of its occurrence in Holocene strata of the
eastern part of the Isthmus lead to the conclusion that the species appeared
relatively late in postglacial time replacing the saiga in Azerbaidzhan.
It probably penetrated directly from the south through the ranges of
northwestern Iran which border the Mugan steppe on the south. This
extension of the range would have been possible in the dry, warm phase of
the Holocene. There is also the possibility that the species was imported
to the Kura lowlands by ancient nomadic tribes which spent the summer
in the northern Iran uplands and the winter on the Mugan steppe.
Cattle herders on the Mil'skaya and Mugan steppes capture young goitered
gazelles which are easily tamed and tether them close to their tents.
In Syria, Palestine and Lebanon and throughout the Near and Middle East
newborn gazelles were caught each year by the thousands for domestication
or for their meat. Usamah ibn Munkidh (1922-23 edition, p. 204) described
the hunting of gazelles: ''When young gazelles are born, the hunters go on
foot and take the young which were born that night, the previous night or
two or three nights before. In one day, 3,000 young were caught near the
Dzhabar fortress.'' Young adult gazelles were sent as gifts to chiefs and
princes, sometimes over very long distances.
The pursuit of goitered gazelles by Median, Albanian, Persian, Arabian
and Mongolian horsemen was probably one of the causes of the separation
of the species range on the plateaus and its penetration into isolated valleys.
Even now when gazelles are hunted on horseback or in automobiles over
long distances, it can be observed that they will seek refuge in solonchak,
tugai and reed thickets or escape into creek valleys or onto the slopes of
clayey ridges. From these places they emerge one or days later onto the
open plain because they are frequently vulnerable to attack by wolves in the
thickets and mountains.
Ancient writings give evidence on the large population and area of
distribution of gcoitered gazelle in eastern and southern Transcaucasia in
the Middle Ages. Nizami Ganjawi (13th century A.D.) frequently employed
the goitered gazelle as a poetic metaphor.
Olearius (1870 edition) mentions herds of goitered gazelles which he
observed in 1633 near Dzhevat [near present-day Sabirabad] on the Mugan
Steppe and commented on the abundance of these animals on the adjacent
Iranian Plateau. In the 18th century goitered gazelle existed on Svyatoi
Island (now Artem Island) which they had reached either through importation
by Arabs for breeding purposes or in a migration from the Apsheron
Peninsula during some postglacial drop in the Caspian Sea level. They were
observed there in 1720-1724 by Soimonov who wrote (1763, р. 110):
"However, the island is inhabited by many wild goats which reproduce there
and it is not known what their food is if not mosses growing on stones."
...''December 3 (1724)... whereas on Svyatoi Island Don Cossacks with
their general killed some wild goats."
Academician Dorn (1875) noted that Svyatoi Island was called
by the Arabs ''the island of steppe зпеер.'' Goitered gazelles were observed
352
357
in the Araks valley in the middle of the last century by Nordman (1840) and
Chopin (1852). They were exterminated in this area only in the second half
of the century through game drives.
The species is adapted to a habitat on level semidesert with a solid
footing (for details, see Vereshchagin, 1937). The animal's great speed
(up to 50-55 km/hr) permits easy escape from wolves and even cheetahs,
except on a boggy soil. It is not found, therefore, on newly plowed or
especially onirrigated soils. Goitered gazelle does inhabit barley and wheat
fields on the Adzhinour Plateau, but only those that are not irrigated and
whose soil is more or less compacted.
While the goitered gazelle is more numerous on level steppes, it is also
encountered in the Tertiary hills of Dzheiran-Chel, Boz-Dag, Turut-Sarudzha
and Kabristan — areas of rugged relief with mountain xerophytes. Deep
frosts and heavy snowfalls bring mass mortality to goitered gazelle
inhabiting the Kura lowlands (Vereshchagin and Dyunin, 1949). In general,
the goitered gazelle is a chinophobe because it cannot reach food under the
snow nor move rapidly over a deep snow cover. This characteristic also
confirms its recent migration to Transcaucasia from the south.
Fluctuations in the Caspian Sea level, breaches of the coastal ridges and
the flooding of the Kura and Araks rivers, as well as changes in the nomadic
and settled populations of eastern Transcaucasia and in their agricultural
techniques, have produced far-reaching changes in the range and population
of goitered gazelle in this area in historical time.
In the 1920's, the distribution area of goitered gazelle coincided with the
saltwort solonchak-semidesert and andropogon steppes, and reached nearly
to Tiflis on the west.
The sharp population decline and the shrinkage of range in eastern
Transcaucasia began in the 1920's and accelerated in the 1930's when the
development of motor and air transport made it possible for poachers to
work mass destruction on the species. It was driven from the Apsheron
Peninsula early in the century and from the Shirak steppe in the 1920's.
In the 1930's hunting with cars and trucks developed to such an extent
that the goitered gazelle was nearly exterminated in the northern part of
the Shirvan steppe. Just in the area around the Kara-Su railroad station
poachers bagged more than 500 goitered gazelles annually. Poachers from
the cotton sovkhozes of the Mil'skaya steppe bagged from 500 to 600 gazelles
each year with the use of automobiles, and this steppe yieldedthe same
number to herdsmen winteringthere. In 1937 goitered gazelle could be
encountered in eight isolated areas of semidesert and poorly-developedtracts
ofthe Kura steppes and the ridges of Adzhinour and Kabristan comprising
approximately 8,219 km?. Its population was in the 5,000-6,000 range
(Vereshchagin, 1939a) (Map 87). During the years that followed, the range
and the number of animals decreased rapidly as poaching techniques were
motorized and as semidesert land was developed for cotton growing.
In the 1940's the species was displaced from the territory north of the
Sumgait River. In Kabristan it was confined to deep semidesert ridges by
the development of the oil industry. Small herds of goitered gazelle could
sometimes be encountered in 1935-1937 on the Gezdek Plateau and near
Mount Kergez 10 km southwest of Baku. By the 1940's only single
Specimens could be observed andthose nocloser than 25-30 km from Baku
in the Kara Dag and Shakhi-Kai region.
353
With the irrigation and plowing of the Kura-Araks lowlands the goitered
gazelle will become extinct in eastern Transcaucasia wu less a steppe
preserve is established and effective measures taken for the protection of
this animal.
Saiga—Saiga tatarica L. Saiga remains are recorded for
Pleistocene and Holocene strata from the British Isles in the west to the
Novosibirskie Islands in the north and Alaska in the east (Nehring, 1890;
Cherskii, 1891; V.I. Gromov, 1948; Pidoplichko, 1951). The stability
of the adaptive features of the saiga from the Middle Pleistocene to the
present points toward a Pliocene or even Miocene origin for the genus.
The lack of findings of closely related forms in Pliocene strata of the
Palaearctic is rather mysterious, but, in view of the adaptation of the
contemporary species, it can be assumed to have developed in steppe and
savannah environments.
Although the Pleistocene fauna of the northern shores of the Mediterranean
generally resembles that of the middle belt of Europe, saiga remains have
not been found in southern France, Italy and Greece (Wolf, 1938, 1939).
The most southerly finds of Pleistocene saiga remains have been in the
Crimean steppes and on the Caucasian Isthmus. On the Russian Plain saiga
bones present massive and certain evidence of Middle and Upper Pleistocene
deposits.
On the Caucasian Isthmus fossil remains of saiga have been reported
from Ciscaucasia and eastern Transcaucasia (Map 86).
There has probably been no penetration of saiga to the south of
Transcaucasia. Nor has it been found in western Transcaucasia which also
has long been a woodless area of rugged relief.
The appearance of the species on the Caucasus can be dated at least
Middle Pleistocene,
358 The saiga remains of at least 70 specimens found in the Binagady
bituminous strata of the Apsheron Peninsula are convincing evidence that
the penetration route of European-Asian steppe forms from the north
followed the western coast of the Caspian Sea. In the pre-Khazar stage
of the ancient Caspian regression, the ecological barrier which the
Khachmas lowland forests of the Kusary sloping plain now present to steppe
species probably did not exist. The winter migrations of saiga recorded
by Glitsch (1865) raise questions on the stability of saiga habitats in eastern
Transcaucasia. Is it possible that these were only seasonal migrations
taken during cold winters? The answer is supplied in a study of the age
groups of animals which perished in the bitumen, based on 60 mandibles
with preserved molars:
1. Young — June-July; milk molars worn down; first true molar erupted:
33750.
2. Young — August-September; first true molar erupted and worn down:
21.7%.
3. Semi-adult — November; second true molar erupted; milk teeth
replaced by permanent teeth: 5.0%.
4. Adults and aged specimens — worn-down true molars: 40%.
This data shows beyond doubt that saiga reproduced on the Apsheron
and inhabited eastern Transcaucasia at least in summer, in fall and at the
start of winter. This habitation continued until some time later, and was
probably terminated only by the Khvalynsk transgression which greatly
constricted the developing steppe belt of the piedmont.
354
359
Saiga inhabited the Trans-Kuban piedmont plain and the Taman Peninsula
of Ciscaucasia in the Upper Pleistocene together with primitive bison and
giant deer. But whether these habitations represented only winter migrations
from the Kuban valley and through the Kerch Strait from Crimea is difficult
to judge from the existing data. The fact that such migrations were possible
is confirmed by the run of saiga on the Aral Sea islands (Berg, 1905).
In postglacial time the saiga range was still enormous, extending from
the Carpathians to Mongolia. Saiga inhabited the Ciscaucasian plain during
the Holocene from the Azov to the Caspian coast. From this area and
particularly from adjacent areas of the Russian Plain multiple findings of
Holocene remains of saiga have been taken. They have been found in cultural
strata in the following proportions (ratio to total number of bones extracted
at each site expressed in percentages): 1) Cossack village of Tsimlyanskaya,
Bronze Age, 15th-10th centuries B.C. — 5% in one excavation area,
12% in another; 2) the farmstead of Krasnyi Yar near Tsimlyanskaya,
Late Bronze Age — 4.2%; 3) the village of Lugovoe in the Assa valley,
Scythian strata — one horn among a great number of remains of domestic
animals; 4) Cossack village of Tsimlyanskaya, Khazar strata, 8th-10th
centuries A.D. — 14.8%; 5) Sarkel fortress near Tsimlyanskaya, 9th-11th
centuries A.D., from 1934 excavations — 6.5%, and from 1950 excavations —
Bee И.
The saiga is not depicted in drawings, sculptures or folklore of the
Caucasian peoples, except for a representation of antelope on a vase from
the Maikop burial (Figure 128) and unrealistic recent drawings that are
almost contemporary reported from Chokrak sandstones near Kapchugai
in Dagestan by Markovin (1953).
In the Ukraine saiga was common in the later Middle Ages (Beauplan,
1823 edition). Litvin (1890 edition, p.48) recorded saiga and roe deer in
the province of Kiev of the Lithuanian Grand Duchy as follows: ''Wild goats
run across from the steppes into the forests in winter and from the forests
to the steppes in Summer in such large numbers that each peasant kills up
to 2,000 of them each уеаг." This was approximately the year 1550.
In the last century the southern range limit of the species in Trans-
caucasia lay alongthe Ust'-Urt, the Aral and the northern Kyzyl-Kum.
During the 19th century and at the beginning of the 20th, the decline in
saiga population оп allsurvivingranges proceeded very rapidly as the result
of direct extermination (Glitsch, 1865; Dinnik, 1914a; and others). The
plowing of land was also a contributing factor. At the time of Rossikov
(1887) saiga still inhabited the lower reaches of the Malka, and in the 1890's
could still be encountered near Stavropol: Dinnik (1914a) reports their
being killed near the Sengileevskoe Lake and the village of Staro-Mar'evka.
At his time saiga were common near Arzgir, Raguli and Turkmenskaya
Stavka [Letnyaya Stavka, Stavropcl Territory] but had become extinct
in the vicinity of Bol'shaya Kugul'ta, Dzhalga, Burukshum and Takhta.
By the beginning of the 20th century they were no longer to be
found in the western Manych area (Bogachev, 1918). At this time numerous
armed detachments on the steppes contributed to the disastrous decrease
in saiga population. They survived only in certain parts of the Chernye
Zemli, in the lower reaches of the Kuma, Volga, Ural and Emba ‘rivers,
in northern Kazakhstan and in Mongolia. They completely disappeared
- from the Terek-Kuma semidesert. In the 1930's migrating herds were
335
seldom seen near Arzgir and Terekli-Mekteb. The establishment of hunting
limits, the confiscation of rifled arms and the gradual depopulation of the
semidesert have brought about-a restoration of the saiga population.
According to Adol'f (1950, 1952) a total of less than 3,500 saigas
inhabited the interfluve of the lower Kuma and Volga in 1939, whereas in
1950 herds of more than 1,500 specimens each were encountered; the total
number of these animals living on the right bank of the Volga was several
tens of thousands at this time.
After World War II saiga again became common on the Terek-Kuma
steppes (Babenyshev, 1948).
In the severe winter of 1948-49 thousands of saiga died on the Astrakhan
and Ciscaucasian steppes from cold and hunger. In the spring of 1950
carcasses of saigas were found on the steppes of the eastern slopes of
Yergeni in groups of five to eight (Kolesnikov, 1950). As snowfalls start
large numbers of saiga rush to the south, toward the Terek and Sulak,
and isolated groups reach the Dagestan foothills. Lavrovskii (1950) reported
the gathering of tens of thousands of saiga north of the Chernye Zemli on
tracts of fully-developed tall weeds.
During blizzards, storms and deep cold in severe winters, saiga, like
goitered gazelle, retreat into the tugai and reeds. Their appearance in
the tugai along the Terek and Sulak is, therefore, not astonishing. When
saiga undertake long migrations, going hundreds of kilometers northwards
in search of moisture to quench their thirst during cold winters without
snow, they are liable to die if they are suddenly caught by a snowstorm.
As a result of such a migration, large numbers of saiga grazed in the corn
and wheat fields of the Grozny Region in 1951.
The present range and habits of saiga characterize it as a steppe animal
which was driven into the desert by man. During the Pleistocene and
Holocene it very probably lived in the forest-steppe and, in any case,
360 inhabited thickets and reedlands of the floodplain, much like some African
antelopes. Long migrations from
forest to steppe and back, even
in winters of unusually light snowfall,
are confirmed by Litvin's cited
report. In general, it may be said
that the present stenotopic features
of the species are only superficial
and, to a considerable degree, forced.
The prospects for the continued
existence of saiga in Ciscaucasia
are poor; plowing and irrigation of
vast steppe and desert areas will
each year restrict further the areas
suitable for the habitation of saiga.
In 1950, 15 saigas were settled on
Bulla Island, south of Baku: they
could probably successfully inhabit
any part of the semidesert of eastern
7 cS м И А,
ААА
с А ААА >
Е |/
FIGURE 144, Representation of a chamois оп а Transcaucasia.
silver bucket from burials in the Trialet ridge Chamois — Rupicapra rupi-
(Kuftin, 1941) Capra’ Cave agtioay Lyd: > Poss
356
361
remains of chamois have frequently been found in Pleistocene and Holocene
cave strata in Belgium, France, Germany and Italy, e.g., the Colomby
grotto which is very rich in animal remains. The finds are almost always
in mountain or piedmont areas.
Only isolated occurrences of chamois have been found in Paleolithic
cave settlements in western Transcaucasia, but the North Ossetian dzuars
contain Middle Holocene remains. None has been found in either Pleistocene
or Holocene strata in the vast piedmont plain region of the Caucasus, nor
have they been encountered in Pleistocene deposits of Iran, Syria, Palestine
and Lebanon. The contemporary distribution lies in the Pyrenees, the
Cantabrian Mountains, the French, Swiss and Italian Alps, the Apennines,
the Carpathians, the Balkan Mountains, Asia Minor and the Caucasus
(Couturier, 1938).
Thus both paleontological data and present zoogeography characterize
the chamois as a mountain species occurring in the alpine fold systems
lying to the north of the western Mediterranean and to the southeast of the
eastern Mediterranean.
Contour representations of chamois, depicting the animal with its chest
pierced by arrows, appear on a silver bucket from Early Bronze Age burials
on the Trialet (Figure 144). Since the chamois does not endure captivity
well and could not have been transported far from its natural habitat by
ancient tribes, this find indicates to archaeologists that the bucket was
fabricated either inthe Armenian Highland or in Asia Minor. The contemporary
range of the chamois on the Caucasus is widespread (Map 88). On the
Main Range, the distribution area can be traced from the west from the
Goitkh Pass and Mount Semeshko (1,032 m) in the upper reaches of the
Pshish (Dinnik, 1896, 1914a; Nasimovich, 1949b), with some occurrences
observed (1955) in northwest Tuapse in the rocks of the Dzhubgi district,
to the east on both sides of the Caucasus as far as Mount Baba-Dag in
Azerbaidzhan. On the western Caucasus the chamois range is from
250-300 m to 3,000 m above sea level. At the end of the last century and
the beginning of the present one the species was very numerous in the
reaches of the Belaya, Urushten and Malaya Laba rivers; as many as
400 chamois could be observed daily and sometimes herds of 90 head were
encountered (Dinnik, 1914a). Between 1926 and 1937, at least 4,000 chamois
inhabited the Caucasian Reservation (Nasimovich, 1949b) and Zharkov
reports as many as 12,000 (1940a). Winter concentrations are likely
to occur on slopes with either a western or an eastern exposure. The animal
is most frequently found in the middle altitudes of the subalpine zone — from
1,500 to 1,700 m.
In the central part of the northern Caucasian mountains, there are
chamois habitations on the slopes of the Elburz and in pine forests of the
ranges which border the upper Baksan, Chegem and Cherek rivers. The
species is not found in the furthest chain of the Skalistyi, appearing only
in Digoria and Ossetia. In Dagestan chamois live among rocks in an area
of subalpine tall grasses and beech—hornbeam forests with a very humid
environment on the ridges of the Chernye Gory to the middle course of the
Argun. Their habitat in Urukh and the upper reaches of the Ardon is
characterized by rocky areas grown with pine forests and stretches to
Nogkau and Tseya. Chamois cannot be observed in central Dagestan which
is deforested, andis found only in small numbers on the eastern slope of
Mount Shakhdag.
857
362
On the southern slopes of the Main Range the species is common in
Abkhazia and Svanetia, and it has found habitats on the very steep slopes
of the upper Kodor, Ingur and Rion gorges in fir, spruce and maple forests,
and particularly in the upper reaches of the Tskhenis-Tskhali, which are
very rich in chamois. There are few to be found in South Ossetia or
Kakhetia, but, further to the east, they are numerous in the beech and
birch forests of the Katekh-Chai and Shin-Chai gorges and in the basin of
the upper Turyan-Chai in Azerbaidzhan.
Herds of 50-70 chamois can be observed in the Zakataly Reservation in
August. In the 1930's the census of chamois in this reservation was
approximately 1,000 head (Markov and Mlokosevich, 1935). The usual
habitats of this animal are the edges of birch forests and very steep sod
slopes grown with matgrass and fescue in those parts of the subalpine
meadow-steppe with bedrock of black slate. On rocky slopes near Kurban-
Efend in the Ismailly region chamois live even in the summer at altitudes
from 800-900 m. In winter, the bulk of the population inhabits beech forests
in the range of 800-1,200 m.
The population and distribution of chamois is considerably less on the
Lesser Caucasus. They are still found along the middle course of the
Chorokh in the Artvin area and, further to the west, on the eastern Taurus,
e.g., the Trebizond vicinity. There are no data on the habitat of chamois
on the northern slopes of the Adzhar-Imeretia Range. At one time chamois
was common in the beech forests and rocks of the Kura left bank in the
Borzhomi area. In 1890 five chamois were penned in on a 130-ha-tract
in the Borzhomi Reservation (Shil'der, 1892). Onthe Trialet ridge, the
species can be observed in the BelyiKlyuch area, although it has been
exterminated in other places. There were probably no chamois in the past
and are none now on the deforested plateaus of Dzhavakhetia.
As late as 1873, Bogdanov and still later Kalishevskii (1904)
noted their presence on Alagez. Neither Burchak-Abramovich in 1944 nor
Dal' in 1950 found any chamois there. In the last century the species
inhabited the northeast margin of the Armenian Highland, and in the 1860's
could be encountered along the Dzegam-Chai and Shamkhor-Chai rivers as
far as the village of Barsun (Radde, 1899, p.74). Folklorist Osipov (1898)
mentions the presence of chamois in the mountainous areas around the
village of Chaikend. By the 1930's it was no longer observable in Karabakh
and on Shakhdag, although Sarkisov (1944g) maintains that it
disappeared from the Murov-Dag only in 1930. Generally speaking,
Sarkisov's data on chamois distribution on the northern slopes of
Shakhdag, on the Lori uplands and in northeastern Karabakh (1944g) require
verification.
The disappearance of chamois from Karabakh and Asia Minor cannot
be explained only by human activity, because on the Murov-Dag and other
ranges there are forests sufficiently deep for the survival of wild goat.
There are no indications of chamois now or in the past in Talysh and
оп the Elburz, although they could certainly exist there. South of the
Armenian Highland, chamois probably exists in relict forest areas. A horn
of chamois from the mountains of the southern coast of Lake Van is
preserved in the Zoological Institute of the Academy of Sciences.
358
363
FIGURE 145. Chamois lying on a scarp in the Katekh-Chai ravine
Photograph by author, 1935
Chamois is less well adapted to the intersecting relief of the mountains
than is the goat. Its skeleton is lighter and its hooves are smaller. It
avoids the rocky places frequented by Caucasian goats (although some
naturalists think otherwise). Its principal enemies are big cats (panthers
and lynx). It can survive in stable rocky areas which are
inhabited by wolf and bear, because it is swifter of movement; it is even
frequently found grazing side by side with bears. The seasonal and daily
migrations of chamois are less pronounced than are those of goats. They
do not migrate in winter to Dagestan where the snow cover is poor, and in
summer they do not leave the forests for the glaciers, because they probably
endure the attacks of bloodsucking diptera better than goats. Their
attachment to biotopes of forest and mesophytic meadow is more stable than
that of goats (Figure 145). On the whole, the chamois is a species adapted
to the mesophytic areas of the mountains rising around the Mediterranean.
It probably did not descend to the plain during the Quaternary because,
being a specialized mountain species but without the speed of the roe deer
or goitered gazelle, it could not move from range to range across the plain
at that time without being destroyed by predators, particularly by wolves.
The absence of chamois remains in plain burials and the data on its
contemporary distribution and adaptations both indicate that the Caucasian
and Alpine chamois derived independently from common ancestors which
existed in the Tertiary on the folded uplands of the Sarmatian sea coasts.
There is, thus, no room for discussion of a chamois migration in the
Quaternary to the European Alps from the mountains of Asia Minor and the
Caucasus or, conversely, from Europe eastwards through the Balkans and
Asia Minor. The Caucasian chamois should be regarded not as a subspecies
of the alpine chamois but as an independent species. A comparison of the
359
two shows the Caucasian chamois to have distinctive morphological features,
including a different voice — a long, hissing sound noted even by
Dinnik (1896).
The absence of chamois from Talysh and El'brus and the rapid decline
of its population and shrinkage of its range in the 19th and 20th centuries
on the Lesser Caucasus indicate that the contemporary ecological optimum
of the species and probably the focus of its origin lie in the ranges
surrounding the Black Sea to the south and east.
The organization of a rational forest and hunting economy requires the
renewed breeding of chamois in Karabakh — in the upper reaches of the
Dzegam, in Shamkhor and in Talysh
West Caucasian and east Caucasian goat— Capra caucasica Guld.
ahd C. cylindricornis Blyth. Fossil remains of representatives of
the subfamily Caprini have been found in Eurasia from the beginning of the
Lower Pliocene, but on the Caucasus the only goat remains that have been
found are from the Pleistocene and Holocene. The Pleistocene localities
are all situated in western and central Transcaucasia on the margins of the
contemporary range, often a distance of 50-60 km in a straight line from
the present habitats of the species (Map 89). The explanation lies partly
in the effects of the glaciations which forced the goats to lower altitudes,
but even more in its widespread distribution in the Pleistocene (see
Chapter II).
To the south remains of local southern forms of C. sinaitica Erenb.
have been found in the Paleolithic of Lebanon, Syria and Palestine and are
recorded under the names of C. beden Wagn., C. ibex L. and
С. cf. nubiana Cuv. (Picard, 1937). Remains of Pleistocene goats are
also reported from many mountain and foothill settlements in Western
Europe — from Ireland and Belgium to Bulgaria. These remains, sometimes
consisting of entire skulls, which are generally related to some species of
the contemporary European goat — C. ibex L., C. aegagrus L.,
С. pyrenaica Schinz. — and to fossil species —Ibex priscus Wold.,
I. cebennarum Gerv. and others, have been found in Pleistocene as
well as in Holocene strata (Woldrich, 1893; Wolf, 1938). A large goat,
Capra sp., inhabited the mountains of the Crimea in the Paleolithic
(Gromova and Gromov, 1937).
364 Carvings of Caucasian goats, especially of the east Caucasian type,
were often produced in the Bronze Age; figures of goats appear on warders,
pendants and seals (Figure 146).
The contour representations on silver vessels from the Maikop burial
show a similarity to west Caucasian goat (Figure 128).
There are no paleontological or archaeological data on an earlier
habitation of the Lesser Caucasus and the Taurus by Caucasian goat.
Nevertheless, it seems very probable that such a habitation occurred,
particularly if one compares Caucasian goat with species endemic to the
area: Caucasian birch mouse, Promethean vole, pine vole, snow vole
and the Caucasian black grouse. The feasibility of Caucasian goat existence
on the Lesser Caucasus is demonstrated in the present day by the
experience of the Borzhomi Reservation (Shil'der, 1892).
The contemporary range of Caucasian goat includes almost all of the
Greater Caucasus for 9 longitudinal degrees (between 39°45' and
48°30'E). The chief habitat is on the slopes of the Main Range, but
360
365
366
they are also encountered on parallel ridges to
the north and south and on spurs with southern
exposures.
In this vast area two contemporary species
developed: the western —C. caucasica
(С. caucasica seve rtzovi) and the eastern —
С. eylindricornis, which differs from the
western chiefly in the structure of the male horns
(see Chapter IV).
The distribution of the west Caucasian goat
(Figure 147) now begins to the west of Mount
Chugush (Nasimovich, 1949a), although in the
i ies 1880's they could be found 30 km westward on the
peaks of Fisht and Oshten (Dinnik, 1914a). In the
FIGURE 146. Kobanian poletop Caucasian Reservation goats inhabit the
of bronze representing east Vodorazdel'nyi Range and its northern spurs.
Caucasian goat — from North On the mountains of Bolshoi Pambak, Dzhuga,
ОБА (ase, Mee) Akhtsarkhva and Yatyrgvarty, the habitats are
somewhat separated in summer (Nasimovich,
1949a). At the start of this century, goats were often encountered on the
Skalistyi ridge and on Mount Achesbok. Temporary habitations by single
specimens have been noted during the last fifty years on the summits of
Dudugush, Chura and Khatsavita.
The west Caucasian goat is also dispersed along the Vodorazdel'nyi
Range and its northern and southern spurs from the upper reaches of Belaya
and Mzymta to the upper reaches of the Balkar Cherek. Very few specimens
are now found in the upper reaches of Teberda and Kodor, but in the
Teberda Reservation, groups of 10-15 can be observed near the glaciers.
Goats have completely disappeared from the upper reaches of the Ingur.
To the east, they inhabit the southern slopes of Tetnuld in the upper reaches
of the Mulkhra and single specimens are seen beyond the village of Ushkul’.
The species is rare on the summits of Shoda and Laila, which are isolated
from the Vodorazdel'nyi Range. There are numerous goats on the northern
slopes of the southern ridges of El'brus. At the beginning of the 20th century,
they lived on Mount Kinzhal, and at the beginning of the 19th century they
were even found in the Pyatigor'e area on Mount Beshtau (Vietinghoff,
1812). Beyond the villages of Tegenekli and Verkhnii Baksan groups of
20-25 goats were observed in the 1950's.
The species is also common in the Bezingi Cherek gorge, where in the
1940's during the fall shepherds killed as many as 40 head. And it is even
more common in the upper reaches of the Balkar Cherek on the slopes of
Dykh-Tau. There in the course of a day in August 1948 the author
encountered three females with young and two young males in the Dykhsu
gorge, and observed numerous tracks and signs of goat-grazing near the
glaciers.
It is very rare in the upper reaches of the Urukh.
The present distribution of east Caucasian goat lies to the east of limits
drawn by the Ingur and Tskhenis-Tskhali headwaters on the southern slopes
and the Baksan and Malka headwaters on the northern slopes. The skull of
a female specimen from Teberda is preserved in the Zoological Institute
of the Academy of Sciences. The Georgian Museum contains in its
361
collections a skull of a male from Teberda with horns of а type showing а
transition from C. cylindricornis to C. caucasica, anda skull
of an 11-year-old male from Baksan showing a transition from
С. caucasica to С. cylindricornis. Thirteen skulls taken from
the northern spurs of El'brus and Mount Kinzhal and now in the Zoological
Institute collection also belong to the east Caucasian form. V.G.Khachvani,
a Svan of Ushkul', near the Ingur source, owns a stuffed specimen, male,
52 years old and typical of the east Caucasian form, which was killed near
the summit of Shkhara in December 1947. From the same village Radde,
in the last century, brought the horns of an aged goat of a transitional
(hybrid?) type between west and east Caucasian.
(365)
FIGURE 147. West Caucasian goats in winter
Watercolor by E.Ya. Zakharov
367
In 1949 Nasimovich (1950) discovered in the Psygansu ravine five skulls
of east Caucasian goat and one skull of west Caucasian goat which had been
discarded by hunters. In finds in the upper reaches of the Bezingi Cherek,
four skulls had horns of the west and central! Caucasian type, and one had
horns of the east Caucasian type. Two skulls found in the Balkar Cherek
region both belonged to the eastern type. Of Dinnik's ten finds of skulls
in 1887 in the Cherkess village of Kundyum two proved to be of the western
form and eight of the eastern (Dinnik, 1890b). All 46 skulls taken from
the Urukh and Ursdon gorges in the villages of Styr-Digor and Nogkau,
which were examined by the author, belonged to the east Caucasian species.
The Rekom dzuar in the Tseya gorge of the Ardon Basin contained
298 pairs of horns, of which 292 belonged to the eastern type and only six
to the western.
From this location the range of the east Caucasian goat is continuous
along the Vodorazdel'nyi Range (Map 89).
According to data collected by the author and V.I. Naniev in North
Ossetia in 1948, the east Caucasian goat can be observed on all the northern
slopes of the Bokovoi and Main ranges and occasionally inthe upper reaches
of the Urukh, Ardon, Fiagdon and Gizel'donrivers. Herds of up to 100 head
are encountered in the upper reaches of the ArkhonRiver and occasionally
in the neighborhood of the villages of Dunta, Arkhon and Aksai.
The species is now rare in the Grozny region and in central Dagestan.
It was probably rare on the Gunib and Khunzakh plateaus even at the time
of the Caucasian wars inthe mid-19th century. Although now completely
extinct in the middle reaches of Kara-Koisu, it undoubtedly could be found
everywhere in that region in the last century. East Caucasian goat is
rarely seen in the Samur and Lakskii areas, but is common in the Avar
Koisu Basin on the Bogos Range, from which Geptner and Formosov (1941)
report herds of 20 head in the 1920's. It is also numerous in the rocky
upper reaches of Kara-Chai and Kusar-Chai on Shalbu-Dag and Shakh-Dag.
There are now few goats on the southern slopes in the upper reaches of
the Aragva and Alazan, and they are completely absent from the Telavi
Range, even though they inhabited this area until quite recently. In the
1930's some 200-300 head were counted in the Lagodekhi Reservation
(Markov, 1940). Following some epizootic outbreak, their numbers were
greatly reduced along the Belokan-Chai and Mazym-Chai (Vereshchagin,
1938a).
The Katekh-Chai ravine is the habitat, both in summer and in winter,
for a particularly heavy population; herds of 200 and more head can be
observed there. The census in the 1930's in the Zakataly Reservation was
3,500-4,000 head (Markov and Mlokosevich, 1935; Vereshchagin, 1938a).
The species is frequently found in cirques in the Shin-Chai and Demir-
Aparan-Chai gorges, and is especially numerous on the southern border
of Dagestan in the upper reaches of the Samur (Figure 148) and in the
vicinity of the three high peaks of Bazar-Dyuzi, Baba-Dag and Shakh-Dag —
a fact noted even in 1896 by Satunin (Radde, 1899). In the 1930's the author
observed there many herds of 30-40 goats, and occasionally aggregations
of 250 or more. The extreme eastern margin of the east Caucasian goat
range passes near Mount Aivasel 25 km east of Mount Baba-Dag.
Today groups of 15-20 goats can be observed near Konakhent in the upper
reaches of the Gil'gin-Chai.
363
FIGURE 148. Khalakhi Lake in the upper reaches of the Samur.
The surrounding rocks are a favorite habitat of east Caucasian goat
Photograph by author, 1937
The eastern Caucasian goat (Figure 149) ranges between wide altitudinal
limits. This animal can be found in summer at altitudes up to 1,200-1,300 m
in ravines of the Zakataly Reservation which are nearly inaccessible from
the south. It frequents mineral springs and passes in valley bottoms at
altitudes of 800-1,000 m. To the east above Vartashen and Ismailly its
summer habitat is on the treeless slopes of the southern ranges at altitudes
of 1,400-1,500 m. In winter the lower limit on snow-covered slopes drops
200-250 та, but remains the same as in Summer in the valley bottoms. After
slaking their thirst atthe mineralsprings, the adult goats either stay near
the riverbed at altitudes of 1,400-1,500 m or climb up to altitudes of
2,800-3,000 m where they rest near snowdrifts, thus completing a daily
vertical migration of 1,500-2,000 m. In August the goats often use passes
at altitudes of 3,400-3,500 m.
On the southern slopes the goat population of the upper reaches of the
Alazan to the upper reaches of the Pirsagat divides in summer, because
the entire alpine meadow belt is occupied by cattle herders with herds of
sheep and cows guarded by dogs. One group of goats spends the entire
summer in the forest, resting on taluses and rock overhangs; the other is
to be found in the passes above the meadow zone on rocks and taluses.
368 The west Caucasian goat of the Recent is now a mesophilous species,
but its adaptation to the humid forest and to alpine meadows is probably
а later, secondary phenomenon.
The contemporary population of Caucasian goat is greater in the eastern,
drier parts of the Caucasus; its preferred pasturage is in meadow-steppe
rather than rich alpine meadows. It is characteristic of the goats of the
364
369
forest populations to take long rests on rocky scarps, taluses and in sunny
forest clearings — a habit which may revert to the ancient ways of their
ancestors whose mountainous habitats were treeless and more arid.
The pattern of Caucasian goat behavior as it relates to firn-snow and
glaciers is more clearly seen in summer than in winter. The animals
generally frequent the firn in August and rest in the sun enjoying the coolness
and the absence of flies (Vereshchagin, 1949a). They avoid frozen slopes
and glaciers, and will look for footing on uncovered stones after a fresh
snowfall. It is rare that they will attempt a crossing over a large glacier
area. During the author's crossing of the huge glaciers near Mount
Laborda east of Mount Shoda in August 1948 no goat tracks were observed
on the glaciers. According to observations of hunters (M.G. Ivkin and
others), large herds of goats migrate each year in November-December
from Lagodekhi and Belokan to Dagestan, always using the snow-free
еее ог 19565.
The general outline of the Caucasian goat ranges has changed little during
the last 150 years. Presumably during the Caucasian wars in the mid-19th
century there were more goats killed than there are now. The population
declined with the establishment of Russian settlements in the Ciscaucasian
piedmonts and the introduction into the mountains of the four-barrelled
Berdan rifle andthe triple-barrelledrifle at the start of this century.
The decline was progressive during the first half of the 20th century
throughout most of the goat ranges. One exception was the hunting preserve
of the Grand Duke in the Kuban Basin where the animals were well-guarded.
The protected environment of the Caucasian and Zakataly reservations
brought about an increase in the goat population in these areas beginning
with the 1930's.
Goats fled from harassment in Fisht and Oshten in the 1880's, and
disappeared from Mount Achishko near Krasnaya Polyana in 1908
(Nasimovich, 1949a). Villagers of Ushkul' killed 200-250 goats in the area
of the Ingur sources over a35—40-year period in the last century. Their
sons, using improved weapons, killed no more than 15-20 in the same time
Span in the first half of this century.
By the beginning of the 20th century, goats had been routed from the
furthest ridges of the Skalistyi in the central part of the northern Caucasus,
particularly from Kinzhal and Barmamyt (Dinnik, 1914a). Dronov collected
thirteen skulls in the upper reaches of the Malka as late as the 1880's.
They have probably been extinct in Pyatigor'e in the Beshtau region since
the beginning of the last century. i
Judging from skulls found in the dzuars of Lesgor and Digorized, the
east Caucasian goat inhabited Digoria early in this century on the Skalistyi
ridge which cuts off the longitudinal valley of the Doniserdon from the
north. Since no fresh skulls have been found in these sanctuaries, the
species must have disappeared by the 1930's-1940's.
The approximate annual yield of Caucasian goat in the 1930's was
probably no less than 4,000 head (Figure 150).
Caucasian goats, then, are not to be considered as on their way to
extinction, even though there is no control over hunting in the mountains.
The existence of three reservations — Caucasian, Teberda and
Zakataly-Lagodekhi — is a guarantee of a continuing, thriving population.
365
The development of cattle breeding, accompanied by the extermination
of wolves and panthers, had both a positive and a negative effect on the
Caucasian goat population. In the early stages of cattle herding, wolves
370 preyed on the domestic cattle in the mountains in the summer and followed
the herds to the plains in winter, thus diminishing the danger to the goat
population. However, as cattle breeding was further developed and pastures
were enlarged, cattle diseases such as scab, foot-and-mouth disease
and others spread to the goat population (Dinnik, 1914a; Vereshchagin,
1938a; Nasimovich, 1941).
FIGURE 149, East Caucasian goats in summer
Watercolor by E. Ya. Zakharov
366
The facility with which Caucasian goats can be crossed with domestic
species, domesticated and acclimatized to the plains, 1еааз фо ап anticipation
of further development of both species through controlled expansion. They
will probably be bred in the Karabakh uplands, on the Shakhdag, and on the
Trialet and the Adzhar-Imeretia ridges, as well as in Talysh and El'brus.
FIGURE 150. Horns of Caucasian goats mounted on a veranda of an Ossetian house in the Urukh gorge
Photograph by author, 1947
The economic effects to be obtained from the hybridization of Caucasian
goats with domestic goats could be considerable (Vereshchagin, 1938a).
Taking into account the endemicity and morphological isolation of
Caucasian goats from other representatives of the genus (see Chapter IV),
the species should be regarded as local in origin, springing from Caucasian
Tertiary ancestors.
Bezoar goat— Capra aegagrus Erxl. Fossil remains of this
species are recorded for Pleistocene and Holocene cave strata in Italy,
Lebanon, Syria and Palestine (Picard, 1937; Wolf, 1938). They have been
found in the Caucasus only in postglacial strata in caves in the Araks gorge
(Polyakov, 1882) and on the Saraibulakh ridge, in Bronze Age strata of the
Teishebaini fortress ruins near Yerevan (Dal', 1940b, 1952), and in dzuars
in Tushetia and Khevsuretia.
371 Bronze Age drawings on limestone rocks in southeast Kabristan west
[? south] of Baku show a similarity to Bezoar goat (Figure 151), and may
indicate a recent habitat for this species on the lower ridges of Kabristan
near the Caspian Sea. The collections of the historical museums of Moscow,
Makhachkala and Grozny contain exquisite bronze figurines of Bezoar goat
372
from Dagestan (Figure 152) which are associated with the cultures of the
Kobanian and Kayakent-Khoro-Chai period (first half of the 1st century B.C. ).
FIGURE 151. Representations of goats on rocks of Beyuk-Dash south of Baku
Photograph by author, 1945
The contemporary distribution of Bezoar goat extends beyond the
Caucasus to the mountains of southwest and central Asia, from Crete to
western India through Turkey, Iran and Afghanistan. There are three
isolated ranges on the Caucasian Isthmus: the eastern part of the Greater
Caucasus and the central and the eastern parts of the Lesser Caucasus.
The upper reaches of the Alazan, the Argun and the Gerdyman-Chai, which
is to say the arid, treeless part of the country, form the limits of the range
on the Greater Caucasus (Map 90). In the 1920's the species inhabited the
Lakskii area of Dagestan, and, according to Dinnik (1914a), could be
encountered 30 km from Gunib at the start of the century.
The Bezoar goat of Dagestan is rarely seen in the forested, humid
slopes of the Main Range between Lagodekhi and Shemakha and in the passes
of the divide — perhaps once in two or three decades.
It is common in the upper Avar Koisu area, in the vicinity of the villages
of Tlyadal, Kidero, Chorod and Takhota, where it inhabits pine and beech
forests. Along the upper Terek and Aragva it is not now observable and,
to judge from the compleie absence of skulls of the species in Ossetian
dzuars, it was probably never an inhabitant of this area.
The range on the Lesser Caucasus is more extensive: it includes on the
southwest the fir forests of the upper Chorokh, certain parts of the
368
373
Adzhar-Imeretia Вапбе, {Ве western spurs of the Trialet in the Borzhomi
District and the southeastern part of the Armenian Highland. The species
is common on the Shakhdag and in the Kedabek and Kalakent areas, e.g.,
in the upper reaches of the Shamkhor
near the Kashkar summit (3,379 m),
on Mount Kyapyaz, beyond Lake
Gek-Gel and below the peak of the
Murov-Dag (3,740 m).
In 1940 a small number of these
goats inhabited the taluses in beech
forests on the southeastern slope of
Mount Kechel-Dag at the eastern
extremity of the Shakhdag ridge.
The guard at the sawmill there killed
seven or eight goats each year.
Mount Kirs (2,743 m) above
Stepanakert was the easternmost summit
inhabited by Bezoar goats in the
1930's.
FIGURE 152. Bronze figurine of Bezoar goat The habitat of this animal in the
MIS Ree Sees) central and southern Armenian Highland
lies in rocky mountains from the Azat
River valley to the Zangezur, Megri and Bargushet ridges. North of the
upland the range is limited by the southern slopes of the Gegam and
Vardeniss ridges (Dal', 1951b). The Bezoar goat is frequently observed
on the Urts and Aiotzdzor [Daralagez] ridges and on the rocky massifs
close by. In 1950 Dal' (1951b) counted 124 goats in five aggregations on
33.5 km? on the Urts ridge.
In the 1870's goats could still be encountered near Elenovka on the Sevan;
Polyakov collected one skull from this region.
In the Nakhichevan A.S.S.R. in 1947 the summits of Alindzha-Dag and
Ilyanlu-Dag in the vicinity of the villages of Abrakunis, Boyanur and
Khanagya provided a habitat isolated by the upland semidesert. During the
historical era, goats were repeatedly routed from these heights by ancient
Armenian tribes which used the summits as natural fortresses. Bezoar
goats are more adaptable than the Caucasian type; they often cross vast
desert areas easily and, therefore, have a great migratory potential.
On the Zangezur ridge and its southern spurs the goat population spreads
eastwards as far as the Araks gorge. It is particularly numerous in the
Negram Mountains between Dzhulfa and Ordubad at altitudes of
1,000-1,200 m. In the Araks gorge it is distributed on the southeastern
slopes of the Megri ridge as far as Okhchi-Chai. In 1947 a small goat
population inhabited desert rocks near Akluis and Ordubad.
There are references in the 14th-century writings of Rashid ad-Din to
the Bezoar goat as an inhabitant of Talysh, where it could still be
encountered in the high mountains, e.g., оп the Kyz-Yurdy summit,
(Radde, 1899, p.75) in the mid-19th century; later it was no longer
observed there.
To the south the range of this goat extends into the dry uplands of Asia
Minor and Iran. Collection specimens were taken from Erzurum, Van,
Kars, Ararat, Tabriz, Sabalan and Damavand at the end of the last century
369
and the beginning of the present one. In the U.S.S.R., the Bezoar goat
also inhabits the Kopet-Dagh and the Bolshie Balkhany. According to
Sarkisov (1944e) it can be observed in Iran throughout the mountains of
Azerbaidzhan, Kurdistan, Kermanshah and Luristan, and to the south in
the provinces of Yazd and Shiraz, in Makran, Baluchistan and as far as
the Sulaiman Range.
The most usual habitat of the species is an arid, rocky mountain biotope
with buckthorn, hawthorn, juniper and other xerophytes. The vertical
distribution of the contemporary Bezoar goat on the Caucasian is from
900 to 3,500 m, but it manages very well at altitudes lower than that, even
down to sea level.
The adaptation of the present range to the eastern Mediterranean area
and the life pattern of the Bezoar goat indicate a protracted evolution
of the species in a dry, hot rocky mountain environment at relatively low
altitudes. The penetration of this goat into the Greater Caucasus probably
proceeded northwards from southwest Asia in one of the xerothermic
periods of the Pliocene or Pleistocene, similarly to that of hamsters of the
genus Mesocricetus. The Surami Range and the steppe-like piedmont
of eastern Ciscaucasia would have provided a likely route. The subsequent
isolation in the arid eastern part of the Greater Caucasus can be explained
by the glaciation of the ranges and the development of a wide mesophytic
zone of forest and meadows to the west. The occurrence of the species in
Dagestan at a later time, e.g., Lower Holocene, is less probable.
Anthropogenic influences on this goat only came into play in the last
decades with the development of cattle herding and the introduction of
long-range guns into the mountains. The decrease in the range is
particularly apparent today on the southeastern Armenian Highland.
In an ideal hunting economy on the Caucasus, the species would have
an unlimited future. Its range could be extended by controlled settlement
along the longitudinal valleys of the central Caucasus, e.g., in the Malka,
Baksan, Cherek, Urukh and Fiagdon ravines where there are places on the
cuesta scarps of the Skalistyi which provide favorable conditions for its
habitation.
Sheep: Argali— Ovis cf. ammon L.—and Armenian mouflon —
Ovis gmelini Blyth. Argali remains are reported from Western Europe
for the Pliocene to the Upper Pleistocene, and mouflon remains for the
Pleistocene to the present.
On the Caucasus occurrences of large sheep of the argali type (Figure 153)
are very rare and have only been found in Middle and Upper Pleistocene
strata of the Transcaucasian mountains and plains. Finds of mouflon
remains are even more rare and have been made only in the Paleolithic
of the Lesser Caucasus and the Mesolithic of Dagestan.
The localities closest to the Caucasus from which finds of Pleistocene
argali have been recorded are caves in the Crimea (Acheulean and
Aurignacian strata). Bones of mouflon-type sheep, O. cf. argaloides
Nehr., were found in Acheulean-Mousterian strata of the Kiik-Koba cave
and in Aurignacian strata of the Adzhi-Koba cave (Gromova, 1935a;
Gromova and Сгоштоу, 1937). In spite of the suitability of the topography
of Palestine, Lebanon and Syria, remains of sheep from Paleolithic and
Neolithic cave strata in these regions are unknown (Picard, 1937).
370
375
FIGURE 153. Remains of argali
1,3 — metacarpal and humeral bones from Middle Pleistocene asphalts near Baku; 2— Mg
from Upper Pleistocene diluvium near Maragheh
A tooth of a large argali was found in Quaternary loams of northwest
Iran near Maragheh, and remains of Ovis sp. were found in caves in the
vicinity of Urmia and Asterabad (Coon, 1951).
There are no occurrences of argali remains in Holocene strata of the
Caucasus.
Aside from the Mesolithic of Dagestan, bones of a small post-Pleistocene
sheep, О. cf. gmelini, have been found only within the limits of the
contemporary range of this species: in the ruins of the Karmir-Blur
fortress near Yerevan (7th-5th centuries B.C.) and in cave strata of the
Saraibulakh ridge (Dal', 1941, 1952) (Map 91).
Depictions of sheep are scant in the ancient literature and representative
art of the Caucasian peoples. The earliest are to be found in sculptures.
A mouflon head carved on a belt buckle from the Trialet burials in southern
Georgia has been related to the second millennium B.C. (Kuftin, 1941).
Bronze pendants in the form of sheep with argali-like horns are quite
common in Kobanian burial and settlement sites. These representations
give an impression, however, of domestic rather than wild animals, as
they do not have the force or vivid quality of untamed animals (Figure 154).
Argali probably disappeared from the Caucasus in the Holocene, not even
surviving in central Dagestan where environmental conditions were most
favorable.
Gold articles have been found in Scythian burials of western Ciscaucasia,
Kellermess and other localities which bear traceries and relief
representations of small-horned sheep of the mouflon type. Similar
representations on so-called ''Sassanid metal'' from Transcaucasia date
from 220-657 A.D. Silver plates in the Hermitage collection show a mounted
hunt of Chosroes I and Shapur II in pursuit of fleeing sheep (Orbeli and
Trever, 1935). The area in which the plates were produced — the Iranian-
Anatolian Plateau — corresponds tothe contemporary rage of southwest
Asian sheep.
371
The present range limit of mouflon-like sheep т the U.S.S.R. is reached
in southern Transcaucasia (Map 91).
Horns and pelts of this species collected from Yerevan, Nakhichevan,
Negram, Dzhulfa, Darry-Dag and the village of Puzian in Zangezur are
preserved in the Georgian Museum. The nearest Iran localities to produce
specimens are Kotur-Dagh, the vicinity of Khoi and the island of Koyun-Dagh
on Lake Urmia.
The collections of the Zoological Institute of the Academy of Sciences
contain horns found by Academician Brosse in Alagez in 1849 and a skull
from the same mountain near the village of Mastart found by Pfizenmair
in 1911. There are also skulls and pelts collected throughout Armenia;
from the Nakhichevan A.S.S.R. (the environs of the village of Shakhbuz,
Dzhamaldin, Arazia), and from Turkey (Mount Ararat, the environs of
Bayazid, the village of Gurdzhi-Bulakh, Pir-Reshid Mountain 60-80 km
from Van Lake). The collections of Mil'kovich (1910), Brandt (1879),
von Vik (1914), Khanykov (1851) and Vvedenskii (1908) contain horns, skulls
and pelts from Kara-Dag to the south of Ordubad, the Zorskie Mountains,
Lake Urmia, Mount Sabalan and the environs of Tabriz.
The distribution of sheep in southern Transcaucasia has diminished
rapidly since the last century. Chopin (1852) comments on the presence
of sheep in Alagaz, but contemporary authors — Sarkisov (1941, 1944a,
1944b) and Ра!" (1945, 1949b) — note that the distribution area ends east
of the Zanga River on the Saraibulakh and Daralagez ridges.
In 1947 the species was rare in
the environs of Abrakunis, Dzhulfa,
Negram and Ordubad and on the Darry-
Dag andIlyanlu-Dag mountains, and
only a small number inhabited the
Zangezur ridge, particularly the
vicinity of the villages of Bichenakh
and Shakhbuz.
The contemporary range of
mouflon in Transcaucasia is
associated with the upland steppes
where groupings of dense xerophytic
grasses and sibbaldia predominate,
with formations on mountain slopes
of thorny astragali in the middle
zone, and with salt—wormwood
groupings in the Araks valley. The
summer habitat of the species is
FIGURE 154. Bronze heads of argali-like sheep in the high mountains and their
from Kobanian burials of North Ossetia descents into the warmer valleys
Uvarova, 1900) 5 Е ;
are made mainly in winter.
It can be supposed from the
general distribution of sheep in Eurasia (Nasonov, 1923) that argali and
mouflon are newcomers to the Caucasian Isthmus. Their range expanded
from a southern focus after the development of a xerophytic landscape.
The distinctiveness of argali-like sheep and mouflon-like sheep cannot
be considered as mere geographic variations of a single species, since the
divergence between the two can be traced as far as the Middle Pleistocene
(Vereshchagin, 1953c).
372
The immediate future of wild sheep on the Caucasus, like that of other
ungulates, will depend mainly upon the effectiveness of hunting regulations.
376 The argali-like sheep should be settled in central Dagestan in Gunib,
Botlikh and Khunzakh.
Primitive and Caucasian bison— Bison sp., B. cf. schoetensacki
Freud., B. priscus Boj., B. bonasus caucasicus Sat. Bison
remains occur in Eurasia from the Upper Pliocene, the earliest finds —
horn stems ofthe smallbison Bisonsp., В. cf. schoetensacki — being
in Upper Pliocene Apsheron clays in Kabristan (eastern Transcaucasia)
and in conglomerates on the Taman Peninsula (Burchak-Abramovich, 1949;
Vereshchagin, 1957a).
FIGURE 155, Caucasian bison
Middle Pleistocene remains of a larger bison — В. priscus
longicornis — апа Upper Pleistocene remains of a smaller one —
B. priscus deminutus —have been found in many localities in
Ciscaucasia and western Transcaucasia. Most of the finds were made in
lacustrine -fluvial and cave strata (Map 92).
The form was rare in the Pleistocene of eastern Transcaucasia and of
the Armenian and Iranian uplands, even being replaced by primitive tur.
Its absence from the Binagady complex is especially puzzling. To the
southwest, however, it occurs quite commonly in caves of Lebanon, Syria
and Palestine in strata from the Acheulean to the Neolithic (Picard, 1937).
The trend toward a smaller size in the bison of Eurasia which can be
observed at the end of the Pleistocene was reinforced and fixed by the
harassment of man and the aridity of the landscape. There is no reason to
attribute this phenomenon to ''deterioration" of the climate in the Upper
373
т
Pleistocene, as 20010515415 and paleontologists have often done. The cooling
of the climate and the development of meadow formations caused an increase
in the size of bison in the Middle Pleistocene. The contemporary Caucasian
bison (Figure 155) is undoubtedly a direct descendant of the local Pliocene -
Pleistocene bison.
At the same time that the bison size diminished in the Upper Pleistocene
and postglacial periods, the populations became isolated in river valleys,
оп high plateaus and in broadleaf forests onthe mountain slopes. The Caucasian
bison has not retained the steppe habits of its Pleistocene ancestors and
American relatives. It has become an inhabitant of the mesophytic forest-
steppe and the mountain forest zone. Its remains have been observed in
certain localities of western Transcaucasia and the Lesser Caucasus.
In the Middle Ages the Caucasian bison population was probably linked
to the bison population of Eastern Europe by way of the wooded valley of
the Don.
The different fates of the Eurasian bison and the American bison in the
Holocene were caused by different anthropogenic effects (Vereshchagin,
1956). The survival by large bison and turs of the destructive mounted
assaults of hordes of Khazars, Polovtsians, Kipchaks and Mongols would
have been strange indeed. They did survive, however, into the 16th century
in Great Lithuania and Mazovia (Vizhener, 1890 edit. ) where they were
hunted with spears ("'metabulo"), and into the 18th century in western Siberia
(Bell, 1776). Pidoplichko's assertion (1951) that the European bison of the
Pleistocene was a permanent inhabitant of the steppe is not well founded.
It is strange that among ancient Caucasian artifacts scuptural and graphic
representations of bison are almost unknown; only some unrealistic
figurines of bulls that somewhat resemble bison have been found among
bronze trinkets of the Kobanians. References to Caucasian bison in literature
date from the Middle Ages.
Rashid ad-Din (1946 edition) writes that when Abaga-Khan was wintering
in Arran in 1275-1276, he and five farsang (Parsees) from Shahrud hunted
the ''mountain buffalo'' in the forest. In describing a hunt of the Ghazan-
Khan in Talysh in 1301-1302 (pp. 188-189), this chronicler reported: ''After
this the warriors organized a round-up and chased the game — mountain
buffalo, dzhurs, wild goats and asses, jackals, foxes, wolves and bears
and all kinds of wild animals and beasts of prey — inside the fence until
they all were gathered in that enclosure. '' The author probably referred to
roe deer ("'dzhuyurs"') as ''dzhurs, '' and to Bezoar goats or goitered
gazelles as ''wild goats."'
It is noteworthy that tales of wild bulls in Talysh and the El'brus
Mountains were repeated up to the 19th century (Ménétries, 1832). Brandt's
critical survey (1867) added nothing new on this subject. It is possible that
the extinction of bison in the forests of El'brus and Talysh occurred at
approximately the same time as it did in the forests of the central
Caucasus, i.e., in the 18th-19th centuries.
The earliest literary reference to bison in Circassia is probably that
of Jean de Luc (1625, 1879 edition) and Arcangelo Lamberti's (1954)
is the first allusion to the species in Abkhazia.
It was known in Moscow that the bison existed in Kabarda, Ossetia and
Ingushetia long before the publications of Academicians Lovich, Guldenstaedt
and others who are considered the earliest discoverers of bison on the
Caucasus (Bashkirov, 1940).
374
378
Page 7994 of the 10th volume of ''The Complete Collection of Statutes of
the Russian Empire, 1649" gives this directive:
"Written Order from the Cabinet of Her Majesty the Empress Anne
Holstein [daughter of Ivan V] to the Astrakhan Obercommandant — on the
yearly catching of various living animals to be sent to the Court and to the
Izmailovo menagerie... It is known to us that there are in Kabarda wild
bulls and: kdosy [bison] which they call dombai, for the sake of which you
shall do your best and not spare money from the Treasury in order that the
local princes shall command the catching of five or ten such bulls and young
cows and send them to the Kizlyar fortress. There they should be lured
with bread, and when they are habituated to it, you shall send them to
Moscow together with the other animals. Write this to the Gori Shah and to
the commandant of the Kizlyar fortress and to the Elmurza of Cherkassia,
asking them to do their utmost for the capture and dispatch of these bulls
and cows."'
FIGURE 156. Bison skulls in the Digorized cave
Photograph by author, 1947
It is possible that bison still inhabited North Ossetia at the time of
Guldenstaedt's journeys in 1770-1774.
Dinnik (1890a) was able to find old men of the village of Zadalesk who told
him how their fathers and grandfathers killed the bison-dombais with iron
bullets. There is an inscription carved on one of the skulls of the Digorized
cave which reads "1833" and may signify the date when one of the last bison
was killed (No.10, North Ossetia Pedagogic Institute) (Figure 156).
3175
By the middle of the 19th century bison survived only in the mountains
and forests of the northeastern Caucasus, from which they gradually
disappeared in several stages. The range limit in the north was probably
the longitudinal course of the Kuban, including the mouth of the Malyi-
Zelenchuk, and in the west the Pshish River. A comparative reading of
recorded folklore led Bashkirov (1940) to assume that bison very possibly
inhabited the upper reaches of the Shakhe, Sochi, Mzymta, Psou, Bzyba,
Kodor and Ingur.
In the 1870's there remained a total of more than 2,000 head of
bison.
Little by little, the Caucasian wars and the settlement of Cossacks in
the foothills drove the bison to the upper Belaya-Laba interfluve. The
decline in population and the shrinkage of the range continued without
interruption as cattle breeding and hunting expanded and deforestation
proceeded.
By the 1890's the Caucasian bison population was estimated at
500-700 head (Satunin, 1898), and by 1918 Kulagin's census was only 442.
379
FIGURE 157. Bison (male) in a fir forest
Photography by Р.Р. Filatov, 1909
The establishment in the 1890's of the Grand Duke's hunting preserve
in a 522-dessiatina tract [1 dessiatina — 2.7 acres] of the Kuban and the
control of poaching in this area halted the decline to a certain extent.
Filatov's investigations (1910, 1912) revealed a further contraction of
the bison range; this map shows a distribution area only in the upper
Belaya-Bolshava Laba interfluve. He placed the bison population at that
time at several hundred, mostly inhabitants of fir forests (Figure 157).
376
After the 1917 Revolution, the Kuban hunting preserve was overrun
by cattle herders, lumbermen, army deserters and hunters armed with
triple-barrelled rifles who destroyed nearly all the Caucasian bison in
that area.
380 In 1919 an epizootic, probably endemic to domestic cattle grazing in the
mountains, broke out among the bison and killed virtually all the rest.
Only 50 animals survived in 1920, at least 20 of them in the Dishi-
Khamyshki district, and these too were subsequently driven off by poachers.
The establishment of the Caucasian Reservation in 1924 could not save
the bison because of the number of small arms among the local population
and the difficulties of protecting the animal in the mountains.
Several bison killings are recorded for the years 1921-1926: near the
Cossack village of Lineinaya, 50 km west of Maikop (1921), near Mount
Gefo (1923), and in the upper reaches of Kodor on Alous and Mastakan
(1925-1926). The latter report from Mount Alous describes the killing
of three bison, probably the only survivors, by Imeretian herdsmen, and
is the last one on record (Bashkirov, 1940).
With the year 1940, the history of the hybrid Caucasian-American
bison begins on the Caucasus (see Chapter VI). These bison possess a
greater vital capacity and can live in the mountain forests throughout the
northern Caucasus and Transcaucasia.
Primitive bull* —Bos trochoceros Meyer, В. mastan-zadei
Bertsch ewe prince mi us (Boy: , By mimutus Ма. Wh 1s"eustomear y.
for a history of the genus Bos to commence with the deposits in northern
India in which many Pliocene and Pleistocene fossils of extinct species of
Bovidae have been preserved.
Like bison, the genus displayed on the Caucasus a successive series of
related forms, which, however, were not as clearly defined as the bison
species. In addition, some species represented lateral phylogenetic
branches.
The history of the primitive European-type Bos — а branch of
B. primigenius —begins on the Caucasian Isthmus with the Middle
Pleistocene.
The most ancient find is the skull of a giant B. trochoceros in
Middle Pleistocene sands of Adzhi-Eilas south of Yerevan (Avakyan, 1946).
Later, Upper Pleistocene finds are recorded for the Apsheron Peninsula
bitumens (Bogachev, 1925b; Burchak-Abramovich, 1951а, 1952d), for the
Armenian Highland and for Ciscaucasia (Map 93). There the history of
Bos onthe Caucasus seems to be broken, although burials of the second
millennium B.C. (Hittite ? culture) on the Trialet ridge near Tsalka and
Kirovokan revealed skulls, metapodia and phalanges of bulls of a size close
to the Holocene wild bull.
Related forms on the Caucasus are those of the Middle Pleistocene
В. mastan-zadei from the Apsheron bitumens (Burchak- Abramovich,
1952d), and of the Holocene В. cf. minutus, а small postglacial bull
from the sands of the Sevan coast (Dal', 1950a). These are isolated
localities. The Caucasian В. mastan-zadei did not become an inhabitant
of forests and did not migrate to the mountains — an interesting point of
* [The Russian text uses "tur" here, but to avoid confusion between bull and goat (since, in English, "tur"
can also mean Caucasian wild goat) we have substituted "primitive bull” in the translation as the common
name for the genus Bos where it is apparent that the reference is to one of the primitive species of Bos.]
Si
381
difference from the bison. The distinguishing ecological characteristics
of these Quaternary species can be observed in the distribution of their
remains in Eastern Europe and on the Caucasus.
For instance, to the north of the Caucasian Isthmus, remains of primitive
bull are well authenticated from alluvial and diluvial deposits of the Russian
Plain river valleys.
The Bos population was less than that of bison throughout its range.
The lower percentage of remains of Bos is particularly apparent to the
southeast. Table 74 shows the breakdown between authenticated bison
and bull skulls collected mainly from river banks and now in museum
collections.
TABLE 74. Distribution of finds of bison and bull skulls on the Russian Plain (data from
museum collections)
Museums
Kuibyshev
Saratov
This distribution confirms to some extent Pidoplichko's statement (1951)
that primitive bulls inhabited more humid landscapes and biotopes than
did bison.
In Europe, the bull was mainly an animal of the mesophytic forest
particularly toward the end of its existence. On the Caucasus and in
southwest Asia this was not the case: it inhabited humid valleys and
meadows of open plateaus and, of course, was easily exterminated. Recent
experience with cattle breeding in the plains of central Asia shows a
parallel: the spring ephemeral growth of the desert and semidesert suffices
for pasturage for only a short time; later, from June onward, grazing is
concentrated in meadow-swamp formations on the shores of lakes that are
not stream-fed (e.g., the Chushka-Kul lakes near the city of Turkestan)
and in the river valleys. Similar feeding habits can be observed in the
coarse-grass steppe: as the dense, scrubby grasses become coarser, the
cattle migrate from the steppe to the ravines, gulleys and humid floodplains.
Attracted as the genus is to meadow formations, the primitive bull
probably found its optimum biotope in the southern regions — southwest
Asia and the Caucasus — at the time of cooling and humidification of the
climate.
The habitation of the Apsheron Bos onthe Pleistocene semidesert —
the eastern Transcaucasian steppes — was probably seasonal, that is,
confined to the period of winter-spring ephemeral growth and not including
the summer. Primitive bulls did not inhabit the forested and swampy
378
Transcaucasus, but, to the south, they were common on the wooded and
steppe -like plateaus of southwest Asia.
Remains of Bos primigenius found in Mesolithic strata of the Belt
cave near Asterabad Bay should probably be correlated with descendants
of B. mastan-zadei, a more xerophilous form than European Bos.
(The same correlation holds for goitered gazelle remains — Coon, 1951.)
В. nomadicus Falc., remains of which are recorded for Lower Neolithic
strata from the site of the ancient town of Annau near Ashkhabad (Duerst,
1908), was probably a descendant of the Middle Pleistocene Apsheron Bos.
Two forms of Bos, one large and one small, probably existed on the
Armenian Highland at the end of the Pleistocene. Judging by the finds of
382 bones of bulls in the Neolithic strata of Urartu, it can be assumed that both
forms were domesticated.
There is no documentation of the existence of Bos on the Caucasus
in our time.
Remains of Bos have been found, extraordinarily enough, in kitchen
middens of ancient towns on the Russian Plain. However, to interpret this
find to mean that the wild Bos was a dangerous enemy of prehistoric man
(Gromova, 1931, p.362) and that man began ''to avoid the hunting of this
dangerous ргеу" when domesticated animals were available is very naive.
We have Shumerian and Assyrian bas-reliefs showing scenes of hunting to
contradict this interpretation.
The explanation lies rather in the sharp decline in the population of this
animal in the Neolithic and in the habits of the European Bos which made
it difficult to hunt. The bison which lived in the forest and was no less
dangerous was hunted throughout the Holocene. In the Middle Ages Bos
were encountered only in isolated localities and were often specially
protected (Vizhener, 1890 edition).
The Bronze Age and, particularly, the Scythian epoch on the Caucasus
have furnished us with representations of bulls: a silver vessel from the
Maikop burial (second millennium B.C.) bears a contour drawing — two
bronze figures are designed in the form of bulls, and flat gold plates show
a bull in profile (Figure 158).
It is likely that contour drawings of bulls and cows on rocks (Figure 159)
in Kabristan south of Baku, probably belonging to Roman time, depict
domesticated animals (Vereshchagin and
Burchak-Abramovich, 1948). A bronze figure
of a powerful bull, similar to the primitive
species, was found in a Kobanian burial in
Ossetia. In the collection of the Historical
Museum of the Academy of Sciences of the
Azerbaidzhan 8.S.R. are many signets and
rings with relief representations of bulls which
date from the first millennium B.C. and were
FIGURE 158. Gold plate depicting found in the vicinity of Mingechaur. There is
a primitive bull from the Maikop a marvellous representation of a bellowing bull
burial (actual size) on a gold signet from the Chertomlyk burial
in the southern Ukraine.
To the south and southwest of the Caucasus, Mesopotamia, Syria and
Egypt produced the wonderful bas-reliefs of the ancient Egyptian, Sumerian,
Babylonian, Assyrian and Hittite cultures, many of them depicting bulls
ВУ)
and bull-hunts, from the second and early first millennia [B.C.] when wild
bulls were common in southwest Asia and northeastern Africa. Usamah ibn
Munkidh hunted bulls in Syria as late as the 12th century.
We could find no reference to the existence or disappearance of Bos
in the histories of the Caucasus. Consequently, we can only infer its history
on the Ciscaucasian steppes and on the plateaus of the Lesser Caucasus
from references to it elsewhere: the ancient Russian chronicles, ''The Lay
of the Host of Igor, '' epic ballads (''byliny'') and ''The Instructions"' of
Vladimir Monomakh (see Dolgikh, 1905, Gromova, 1931, Kolesnik, 1936).
It is very probable that Bos became extinct on the forest-steppe of
Ciscaucasia and on the Lesser Caucasus some time earlier than it did in
Europe.
383 The difficulties of evaluating paleontological, zoogeographical and
ecological data to determine species origin and fauna formation are
significantly revealed in this survey of the appearance, development and
contemporary status of certain Quaternary mammals of the Caucasus.
These difficulties are all the greater in elucidating the origins of
eurytopic ubiquitous species (such as European hedgehog, fox and wolf)
and several other mass species of the Quaternary (such as deer and bison)
which have nearly stable distribution areas. Nevertheless, it is apparent
that the local nucleus of the Caucasian mammalian fauna of the Tertiary
formed in the Pliocene or even in the Upper Miocene, whereas for the
most part the ancestral forms of Quaternary species do not occur in the
strata of these periods.
FIGURE 159, Representation of bulls on the rocks of Beyuk-Dash south of Baku
Photograph by author, 1945
1704 380
384
The development of the ranges of many Caucasian mammals was
demonstrably dependent upon the formation of Quaternary topography.
The characteristics of the ranges depend to a lesser extent upon the
species development of morphological-physiological adaptations and the
interrelationships of biocenoses.
The history of the development and disappearance of herbivore ranges
even suggests that the processes proceeded autonomously within each
species, each independent of the other; the harassment of predators exerted
a secondary influence on herbivore distribution.
Three principal types of ranges characteristic for Caucasian mammals
can be distinguished from species distribution and ecological
interrelationships:
1. Caucasian forest and alpine type, associated with Tertiary mesophytic
forest and mountain-meadow landscapes. Examples: the ranges of moles,
Promethean vole, pine vole and chamois.
2. Southwest Asian mountain-desert and mountain-steppe type,
associated with Tertiary xerophytic landscapes which developed on the
Caucasian landmass in the arid, hot Cezonoic period. Examples: the
ranges of Persian and red-tailed gerbil, Asia Minor hamster, steppe vole
and wild goat.
3. European-Asian steppe type, associated with steppe landscapes which
developed on the Ciscaucasian plains in the Pleistocene. Examples: the
ranges of corsac fox, little suslik and saiga.
The secondary phase of mammalian development proceeded during the
Quaternary from the bases of these three ranges and resulted from the
settlement in the Pleistocene and Holocene of European, central Asian and
southern Asian species on the Caucasus. The range of central Asian desert
species is the most isolated and alien to the Caucasus (see Chapter V).
Data drawn from the Recent and based on paleontological studies of the
distribution areas and on observations of the ecology of the species under
discussion provide a broad chronological scheme of the origins of
Caucasian Quaternary mammals. Three different age (stratigraphic) groups
can be isolated in this fauna: Tertiary (Pliocene), Pleistocene and Holocene.
The assemblage of local Tertiary forms (or their ancestors) which
undoubtedly still existed on the Caucasus in the Pliocene was composed of:
Insectivora, Chiroptera, Carnivora, Rodentia, Proboscidea, Perissodactyla
and Artiodactyla — these formed a part of the base of the Holocene fauna.
The assemblage is heterogeneous in both ecology and origin and is subdivided
into mesophilous and xerophilous classifications.
The species of the first — mesophilous — subgroup are local; they
appeared in mesophytic eastern Mediterranean landscapes, i.e., mountain
and lowland broadleaf forests and alpine meadows. They are the Caucasian
mole, trogontherium beaver, European beaver, black rat, Promethean
vole, Caucasian snow vole, long-tailed snow vole, roe deer and Caucasian
goat.
The species in this group are eurytopic and ubiquitous. Pleistocene
deposits contain remains of related forms: white-toothed shrew, fox, wolf,
bear, deer, tur and bison. From this group the principal survivors are
the ubiquitous species, the mountain-forest species and those other species
which could adapt to the mountain-forest.
381
385
The second — xerophilous — subgroup is composed of thermophilous
species, endemic to arid, hot habitats of the eastern Mediterranean,
known in zoogeographical literature as Iran-Asia Minor and Mediterranean
species. Most of the species migrated to the Caucasian Isthmus from the
south and their principal ranges lie in the southern and southeastern
Caucasus and extend to the south of the Caucasus. The others survived
on the Caucasus from the Pliocene, e.g., the Georgian macaca.
Typical of this subgroup are the stone marten, Asia Minor suslik,
common hamster, Persian gerbil, steppe vole and wild goat. Some of these
forms can be found now in xerophytic locations on the Greater Caucasus
and its foothills, and some only on the southern borders of Transcaucasia.
It is very probable that this subgroup comprises relatively eurytopic
animals, which, however, tend to be more xerophilous and southern in their
adaptations. Examples are the long-eared hedgehog, tiger polecat
and migratory hamster; the range of the latter passes through the
eastern half of the Caucasian Isthmus, extending from the mountain steppes
and semideserts of northern Iran to the steppes of the Russian Plain
(see Maps 1, 26, 60). The ranges of some species in this subgroup were
developed on the Caucasus in the Pleistocene, in some cases to a greater
extent than in the Holocene, e.g., the common hamster and other southwest
Asian forms. In other cases, the ranges developed in a northerly direction,
i.e., onthe plains of Transcaucasia and even in Ciscaucasia in the Upper
Pleistocene and as recently as the Holocene, e.g., Asia Minor gerbil,
kulan and goitered gazelle.
European-Asian hydrophilous and forest species formed the fauna of
the Isthmus. The emergence of European forest-type animals on the
Caucasus from Western Europe was feasible in the Tertiary, the Pleistocene
and all epochs when the broadleaf forests of Europe and the Caucasus
were connected through Asia Minor and the Balkans or through the Russian
Plain. The formation of parallel species, especially in mountain forms
like lynx, Caucasian snow vole, roe deer and chamois, was also possible.
Those species which penetrated the Isthmus from the north, southwest
and southeast belong to the younger — Pleistocene — genetic group. The
reasons for their settlement lie in the ancient formation of the landmass
of the northern Caucasian Isthmus, in the development of forest and
meadow-steppe landscapes during the time of climate-cooling, in the
development of steppe and semidesert landscapes in the Ciscaucasian plains
during the time of climate-warming. It is very probable that there were
two migratory routes to and from the Caucasus open to forest species in
the Pleistocene — a northern one along the Don floodplain, and a southwestern
one along the northern coast of Asia Minor.
By either of these ways, the elk and the typical European brown bear
could have penetrated the Caucasus. Only the northern route would have
been feasible for the striped field mouse, the large Asian roe deer and
other species; pine marten and European wildcat are more likely and
common red-backed vole is certain to have followed the southwestern route.
The time of their emergence on the Caucasus probably varied for each of
these species — common hamster penetrating a little earlier, European
brown bear and Asian roe deer a little later.
The Pleistocene assemblage also included steppe elements: corsac fox,
little suslik, saiga and other species which emerged on the Caucasus in
382
386
the Lower Pleistocene, and steppe lemming which emerged later. In the
Pleistocene, the northern steppe influence reached even into eastern
Transcaucasia.
The development of southern southwest Asian species was difficult in
the Pleistocene because of the progressive cooling.
The so-called interglacial epochs, warm, dry periods, can be traced
on the Isthmus from the distribution areas of Turan desert species and,
to some extent, of southwest Asian species.
The latest genetic group to emerge on the Isthmus in the Holocene was
composed of southern, thermophilous species which migrated to the
Caucasus during the postglacial warming and dessication of landscapes.
It is comprised of jackal, striped hyena, lion, tiger, jungle cat, kulan,
goitered gazelle and probably some Insectivora and Chiroptera. Some of
these species became extinct on the Caucasus as the result of human
activity.
The Norway rat and the European races of house mouse can be counted
as casual species, which were brought by ocean and river vessels of the
ancient Greeks, Slavs, Khazars and Varangians, and later by railroad
and automobile.
A study of the characteristics of former and contemporary ranges of
Caucasian mammals shows the effects of a great natural dynamic on the
populations and on the range patterns of isolated species during the
Quaternary. It also reveals the significant relationship between the
anthropogenic influences of the historical epoch and the populations and
ranges of many species.
The discontinuities in the ranges of stenotopic and relatively stenotopic
mammals of the Caucasus can be explained in the majority of cases by the
alternation of dry and humid epochs during the Cenozoic and the orogeny
and glaciation of the mountain ranges.
The most characteristic are the high-mountain breaches in the ranges
of Caucasian endemics — Caucasian snow vole, Promethean vole and
chamois — in the region of the Surami Range; their age is probably Lower
Holocene. Discontinuities in the ranges of mesophilous and forest species,
e.g., mole, pine vole and roe deer, are peculiar to central Ciscaucasia
and eastern Transcaucasia; some date from the Pleistocene and some from
the post-Glacial. Among the xerophilous species, breaks can be observed
in the ranges of common hamster, steppe vole and wild goat.
An analysis of discontinuities like these, particularly those occurring
in the ranges of rodents, reveals the patterns of two xerothermic epochs
on the Caucasus — one in very ancient and the other in Recent time. The
effects of a cold, humid epoch between these two can also be traced.
Chapters IV and V are devoted to a general analysis of the distribution
of all Holocene animals, based on zoogeographical maps, and to an attempt
to evaluate with some precision the degree and kind of faunistic influences
of the adjacent territories through a study of the stratigraphic and
geographic variabilities.
383
387
388
Chapter IV
STRATIGRAPHIC AND GEOGRAPHIC VARIATION*
IN CAUCASIAN QUATERNARY MAMMALS
The study of the phylogeny of Caucasian Quaternary mammals presents
many problems.
The fossil remains from most localities are fragmentary, and suffer
from post mortem crushing, making biometric studies very difficult. Skull
material and skins of Recent small animals have been collected in different
years and seasons by many workers using a variety of preservation. For
these reasons conclusions on the spatial distribution and evolution of animal
Species are not always reliable and convincing. For example, the annual
and seasonal changes in growth rate and weight among rodent populations
with short life cycles (e. g., moles and voles) may be more pronounced
than variation which is related to local environmental changes.
Insectivores and murids preserved in spirit and by dry method are of
little use for detailed systematic study, as pointed out by Shidlovskii
(1953a,b, 1954a, b). Data on geographic variation in fur-bearing animals
is mostly based on information from the fur-trading stations and must be
accepted with reservations, as a study of this type must take into account
individual, sex and age variability within the populations.
It is felt that comparative study of the morphology and systematics of
the mammals of the Caucasus and the Russian Plain may contribute to an
understanding of the development of the fauna, and therefore a summary
of data on some orders is presented below. Most of the work on the evolution of
the species through time has been based on better material, more amenable
to biometric study.
Order INSECTIVORA
The number of fossil insectivores in the Caucasus is negligible. The
Middle Pleistocene hedgehogs (He miechinus aff. auritus and
Erinaceus aff. europaeus) from the bitumens at Binagady on the
Apsheron Peninsula are practically indistinguishable from the Recent forms
in size and structure of the mandibles and teeth.
The geographic variation of the common hedgehog in the Caucasus is
reflected in the darker spines and abdominal fur in gray woodland and
wooded lowland populations. Dark varieties —Erinaceus europaeus
concolor, E. europaeus ponticus —occur onthe Black Sea
* Because it seems to us more correct to refer to evolutionary and geographic changes in living organisms as
variation we reserve the more commonly used term variability to denote individual and age changes.
384
coast of the Caucasus and in Asia Minor, the Eastern European
EF. europaeus rumanicus is found in,Ciscaucasia;) and the light-
colored Е. europaeus transcaucasicus in the semideserts of
Transcaucasia.
The variation in the Recent long-eared hedgehog is slight over the
Caucasian Isthmus. The populations of the small forms, Hemiechinus
auritus calligoni and H. auritus brachyotis, occur in the
semideserts of eastern Ciscaucasia and in the middle Araks valley.
Detailed information on the stratigraphic variation of the Caucasian
moles (genus Talpa) is not available. The Recent mole populations
can be subdivided into distinct geographic varieties, particularly in the
areas which became isolated long ago.
The biggest Caucasian moles (Talpa caucasica ognevi) inhabit
the southern slopes of the western half of the Greater Caucasus and the
Trialet ridge (Ognev, 1926a, 1928; Stroganov, 1948)*. The moles decrease
in size to the south and southeast. Populations of the smallest varieties,
т. Opeee vanieay) Sh Glamsye-ayicays са. and В. on ile mctiad Ws
talyschensis, occur in the ridges of the Armenian Highlands and in the
forests of Talysh. These populations are isolated from the main area of
distribution of the species; however, N.K. Deparma has recently found
a relatively small population of moles in the southeastern Caucasus, even
smaller than the Talysh population.
The size of the skulls and metatarsals of the Caucasian moles is given
in Table 75.
(385) тАвьЕ 78. Geographic variation in size (mm) of Caucasian motes*
Measurements
Lesser Caucasus Talpa
orientalis trans-
caucasica ognevi
caucasica
West of Greater and
Lesser Caucasus Talpa
8 specimens
Greater Caucasus
Talpa caucasica
caucasica
Lesser Caucasus
Talpa orientalis
orientalis
170 specimens
Talpa orientalis
talyschensis
3 specimens
240 specimens
62 specimens
Talysh
29.6
35.9 33.9 30.1 32.4
Condylobasal length
Крео ее З5.1-317.2 33.1-395.7 29.8-32.6 31.3-33.0 29.5-29.8
Length of upper tooth 14.5 13.4 IDE 11.2 11.1
ее оо 14.0-15.0 12.5-13.8 11.9-12.7 10.5-11.7 11.0-11.2
19.6 18.3 16.9 16.5 15.5
о. 190-200 17.5-19.2 16.0-17.9 14.2- 17.5 15.0-16.0
Note. Mean value of observed ranges in the numerator, limits in the denominator.
* Measurements by Stroganov (1948), Dal’ (1944a), Vereshchagin (1945b) (Talysh).
This size decrease to the southeast may be attributed to the prolonged
isolation of moles on mountain ranges under conditions of increasing
* In 1948 S.U. Stroganov tentatively identified 8 specimens of moles from the region of Kutaisi and
Borzhomi as the Italian mole T. romana ognevi. Subject to the final clarification of affinities
of the Caucasian and west Mediterranean moles, we regard this form as a subspecies of T. caucasica;
T. europaea transcaucasica, described by Dal' (1944a) from the Pambak ridge, is tentatively
referred to T. orientalis.
385
draught, increasing annual temperatures and depletion of food resources,
i.e., land invertebrates.
The speciation of the Caucasian shrews can be understood in the light
of the fact that these mesophilous forms, like the moles, were isolated for
a long period in the forest-mountain massifs.
Recent shrews of the genus Sorex onthe Caucasian Isthmus (e.g.,
Sorex minutus) also decrease in size from west to southeast. The
Caucasian population of the common shrew is at present regarded as a
subspecies of the European shrew, S. araneus satunini (Ognev, 1928).
This shrew is distinguished by a short, wide nasal region and shorter
mandible, etc.
There are, however, certain larger local species of shrews which have
evolved in the Caucasus, for example, the darker-colored and long-tailed
S. raddei, closely related to the European S. araneus.
European water shrews in the Caucasus (genus Neomys) are
morphologically and ecologically more specialized than land shrews, but
their genetic relationships are not well known. The forms described —
Neomys leptodactylus, N. schelkovnikovi, N. balcaricus
and №. dagestanicus —are regarded here as subspecies of the European
N. fodiens. The first two of the above are characterized by narrow
paws, as pointed out by Satunin (1915a), whereas the Dagestan
subspecies is characterized by a poorly-developed tail, pale coloration, etc.,
characters typical of animals inhabiting lands with progressive desiccation
(Spain, the Crimea).
The structure of the skull and teeth of the white-toothed shrews of the
Middle Pleistocene from the Binagady burial (Crocidula russula,
C. leucodon) does not differ significantly from the contemporary forms
inhabiting eastern Transcaucasia. Southern white-toothed shrews are much
more widely distributed over the Caucasian Isthmus than other species;
particularly abundantis C. russula guldenstaedti which inhabits
both the dry and the humid zones.
Geographic color variation is well seen in the Recent species and is
directly related to the aquatic biotopes in the hot semidesert and humid
shady forests of Colchis and Asterabad. According to Shidlovskii (1953b),
the long-tailed shrews of Transcaucasia are smaller and darker in the
humid forests of the coastal plains of the Black and Caspian seas than inthe
390 center of the country and inthe dry areas of eastern Transcaucasia (Table 76).
Clearly, if the isolation of the Caspian shrew can be proved, then any
similarity in color will be the result of convergence due to the ecological
similarity in the areas of Colchis and Asterabad.
389
Order CHIROPTERA
With a few exceptions, fossil Chiroptera are known in the Caucasus only
from the Holocene beds. Thus there is no basis for discussion of the
phylogenetic changes in this order. According to the data available in the
literature (Bobrinskii, Kuznetsov, Kuzyakin, 1944; Kuzyakin, 1950), the
geographic variation in some Caucasian Rhinolophidae is expressed in the
development of lighter coloration from northwest to southeast and in the
increase in body size with increasing altitude above sea level
(В. hipposideros, В. ferrum equinum).
386
391
The widely distributed genera Myotis and Vespertilio are
sometimes intermediate between the European-Siberian and central Asian
populations (e.g., Myotis mystacinus), while other species (e.g.,
Vespertilio se rotinus) are closer to the European forms.
TABLE 76. Geographic variation in dimensions (in mm) and coloration of long-tailed shrew in Transcaucasia*
Western Transcaucasia, Central Transcaucasia, Eastern Transcaucasia,
Colchis Kura valley Talysh
Dimensions and color Crocidura russula Crocidura russula Crocidura russula
monacha guldenstaedti caspica
150 specimens 170 specimens 40 specimens
70.6 74.1 70.6
Body length......... 60-89 60-91 63-80
12.1 13.0 13.1
Foomlength о бою 69 9 6 11.0-15.0 11.0-15.0 12.0-13.6
Condylobasal length of 18.1 18.7 19.3
Stila teateete ciel -wentelfeiil« 16.1-19.4 17.5-19.8 18.6-20.2
Length of upper tooth 8.1 8.2 8.9
FROM ооо, Ome ba Choo 7.6-8.8 7.3-8.9 8.3-9.4
Whole body dark
chestnut brown, tail
very dark all over
Upper part of body gray-
brown, lower part
grayish white; tail
uniformly dark
Whole body dark chestnut
brown, tail uniformly
dark
Note. Mean value of observed ranges in the numerator, limits in the denominator,
* According to Shidlovskii (1953b), whose measurements are given in the above table, C. russu la
caspica must be identified as a distinct species, C. caspica. This, however, needs further study,
since the long-tailed shrew is a highly variable species, both ecologically and morphologically.
In order to reach reliable conclusions on the geographic variation in the
Caucasian bats, good southwest Asian data must be studied, as the ranges
of distribution of most of the species extend from the Caucasus far to the
south and southwest.
Order CARNIVORA
The paucity of carnivore remains in Quaternary burials causes difficulties
in the study of the morphological evolution of this order. The only exception
is the Binagady locality, where there are particularly numerous canid
remains.
The Middle Pleistocene Caucasian wolves are characterized by a smaller
brain cavity, narrower nasal cavities, a narrower cheek region, shorter
upper tooth row and smaller intertemporal width (Figure 160). It is of
interest to note that no differences were recorded in the shape and
structure of the teeth themselves. The decrease in the length of the upper
tooth row was due both to closer spacing of the teeth and to the gradual
disappearance of M*. In 33 wolf skulls from Binagady, M® was present
387
392
Frequency
=~ MH WH HN DBS DW &
2 16 20 24 28 32 36 40 44%
FIGURE 160. Variation in the temporal index (ratio
of the intertemporal width to the length of the brain
case expressed as a %) of Middle Pleistocene wolves
from the Caucasus
1— Recent Canis lupus cubanensis; 2—
Fossil C. lupus binagadenis, eastern
Transcaucasia, Binagady. Dots indicate mean
values
in only 2 (i.e., 6%). Out of 222
skulls of Recent wolves from the
U.S.S.R., МЗ was found in only
0.9%. Changes in the limb bones
are reflected in the gradual
narrowing of the scapulae with time
(Vereshchagin, 1951b).
The present geographic variation
of the wolves on the Isthmus has
been poorly studied; however, it is
known that the wolves of the
Transcaucasian plains are somewhat
smaller than those of Ciscaucasia,
while the Armenian Highland forms
are larger. According to Dal'
(1951a), the Armenian Canis
lupus hajastanicus is no
smaller in size than the Kuban wolf,
and is distinguishable from it by its
long, soft fur; its back is bright
whitish gray and yellowish with a
black tint. The fur traders in the
Caucasus distinguish the ''steppe"’,
relatively light-colored wolves of
the open plains, and the darker
varieties from the ''forest''. Wolves
of bright iron-rust color occur in
Kabarda. This is a fairly rare
example of an aberrant variety.
Three skins of this variety are in
the collections of the ZIN Museum.
The probable Middle Pleistocene ancestor of the Recent fox, Vulpes
khomenkoi from the fossiliferous beds in eastern Transcaucasia, is
characterized by small molars and short canines, like those of the Arctic
fox (Bogachev, 1938c; Vereshchagin, 1951b). Towards the end of Middle
Pleistocene times (Khazar stage) the fox evolved into the modern form and
the subsequent changes in the structure of the skull have been negligible.
The cranial features of the Middle Pleistocene foxes of eastern
Transcaucasia (Binagady), however, are quite variable. This variation is
characteristic both of the Recent fox of the Kura Lowlands (Vulpes
vulpes alpherakyi) and the foxes of the highlands — У. vulpes
kurdistanica, V. vulpes alticola. The basic structure and size
of their teeth, however, does not differ from the former species, as shown
in the graphs (Figure 161).
The skull size and fur of the Recent Caucasian foxes display a marked
geographic variation. The general pattern of variation has been traced on
individual skulls and pelts by Ognev (1931), who published a distribution
map of the described subspecies of Caucasian foxes: Vulpes vulpes
stepensis, V. vulpes karagan, V. vulpes caucasica,
Vevilpes aitreo ra,
alpherakyi.
V. vulpes kurdistanica, V. vulpes
393
(392)
The distribution of the Caucasian foxes, based оп the differences in the
fur coloration (material studied at fur-trading posts), has been accurately
described by Kuznetsov (Bobrinskii, Kuznetsov, and Kuzyakin, 1944).
The first distribution map of local variation (in such features as fur
color and other associated characters) and size variation in the Caucasian
foxes was published by us (Vereshchagin, 1947d).
Pelts obtained in the course of hundreds of years are subdivided into
six races by the local fur traders: Ukrainian, Don, Kuban, North Caucasian,
Yerevan and Transcaucasian.
Our study of the pelts of more than 13,000 specimens of foxes at the
Rostov and Tbilisi fur stations may be briefly summarized as follows.
Ukrainian. This race is subdivided into north and central Ukrainian.
These are big foxes with relatively stiff guard hair, rusty white on the
back and straw on the sides. The belly is mostly rust in color and
the upper surface of the paws is black. This variety occurs, though rarely,
in the Ciscaucasian plains. Populations of these foxes occur in pockets
among the typical north Caucasian foxes, from the northwestern Manych
region to the Kabarda plains. According to the fur traders, the presence
of this variety in the 1940's in Ciscaucasia was noticed only after World War II.
Don. Fairly large light-colored foxes, rusty white, with red shoulders
and belly. The guard hair is shorter than in the Ukrainian race. The most
commonly occurring colors are probably related to age and sex: rust, light
rust, rust gray; more rare are dark rust and gray rust specimens.
These foxes lived round the coast of the Sea of Azov, the lower reaches
of the Don, and the Sal'sk steppes. In the east their range extends to the
Yegorlyk, in the south, to the Yeya.
ш
г
Ss
2
—— oe a ee ===.
Frequency
~ % &S RH @® NN @ >
B>
\
|
|
}
}
|
|
|
|
|
\
< № &® BA FW HN & <
56 58 60 62 64 66 68 3056 9501 122 13mm
FIGURE 161. Variation in tooth size of Transcaucasian Middle Pleistocene foxes
I — length of lower molar row; II — length of crown of My; Ш — length of crowns of Mg+M3;
1 — Recent Vulpes vulpes alpherakyi; 2— fossil V. vulpes aff. alpherakyi;
eastern Transcaucasia, Binagady. Dots indicate the mean values
389
394
Kuban. These foxes are somewhat larger in size than the Don race.
The characteristic bright rust and even reddish color on the face, back
and, particularly, neck and shoulders produces a noticeable ''cross"'
pattern. The guard hair on the back is shorter than that of the Ukrainian
race. Individuals of very rich rust color are most common, while rust
gray individuals are less abundant, and even more rare are the light rust
forms.
The range of this race is on the steppes of the Kuban region, on the
Trans-Kuban inclined plain from the Kerch Strait in the west to the Kuban-
Kuma water divide in the east.
North Caucasian. These are foxes of a somewhat smaller size than those
of the Kuban race. The fur is coarse; on the back the guard hair often
forms ''curls'' — patches of standing coarse hair. The belly is black to
dark gray. These foxes occur in two varieties, the red and the paler red-
gray forms, the latter becoming more abundant to the southeast. The
foxes identified аз У. vulpes caucasica belong to this variety.
The distribution of this race is from central and eastern Ciscaucasia,
from the Trans-Kuban Plainto the Caspian coast, and from the lower Kuma
and Terek to the Samur. Pelts of such foxes also come from eastern
Transcaucaia and show great similarity in coloration and other features
with the eastern Ciscaucasian form,
Transcaucasian. This race is represented by the small foxes of the
eastern Transcaucasian plains. They are subdivided into three varieties:
Transcaucasian red, Transcaucasian red-gray, and Transcaucasian gray.
Their fur is relatively coarse and short, often with considerable admixture
of black hair on the rump and shoulders. The foxes are characterized by
a yellowish rusty "Бапа" of variable width, which runs along the length of
the back. The band is particularly conspicuous in the paler colored young
specimens; as arule, young individuals are gray in color. The small fox
of eastern Transcaucasia, described as V. vulpes alpherakyi,
belongs to this race.
Foxes of the Transcaucasian race inhabit the lower reaches of the Terek
and Sulak, the western coast of the Caspian, the plains of eastern
Transcaucasia andthe middle Araks valley. Inthe forties, the Transcaucasian
Vulpes vulpes karagan, disseminatedinthe fox population of eastern
Transcaucasia, was identified as belonging to this race.
By the state standards of the forties, this is a small fox with soft
uniformly dark gray and even somewhat dark brown fur. The limbs below
the elbow and knee joints are black or black-brown. This variety has little
in common with the central Asian karagan fox, as the former is an
aberrant form, а dark-colored variety of У. alpherakyi, like the black-
brown varieties of the arctic fox.
Yerevan. A large fox, though smaller than the Ciscaucasian fox, with
a weakly ossified skull. It is clearly distinguishable from other Caucasian
varieties by its extremely thick and soft fur. The guard hair is very long
and silky to the touch, and the color varies from light yellow to a dirty
rust brown. Foxes identified as V. vulpes alticola and V. vulpes
kurdistanica belong to this race,
390
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391
(396) TABLE 78. Geographic variation in size (cm) of fresh fox pelts from the Caucasian Isthmus and the Ukraine
Length from
nose to tail
base
Number of
skins measured
Area of
pelt (cm?)
Middle
width
Length of tail
Races and fur station Е
without fur
Central Ukrainian, Stalino
Don, Novocherkassk ..... 34
Kuban, Slavyanskaya ... 30
North Caucasian, Blagodarnoe 28
Mongrel, Batumi 28
Mongrel, Tsalka 34
Transcaucasian, Yevlakh 36
Transcaucasian, Baku . pe
Yerevan, Теа ан 23
Уетеуап,, Yerevan.) 25190
Note. Mean value in the numerator, observed range in the denominator.
These species inhabit the Armenian highlands, and probably the ranges
of northwestern Iran and eastern Turkey in the south. Similar though not
identical skins come from the highlands of the Greater Caucasus,
particularly from south Ossetia and central Dagestan.
It should be mentioned that the age variation of the animals could not be
taken into account in the construction of the distribution map of the fox
races (Figure 162) and the diagram of the distribution of different color
types (Figure 163). The distribution and relative proportions of the different
races are given in Table 77.
307 Unusual coloring is observed occasionally in certain strains of Caucasian
foxes, e.g., the ''sivodushka'', which has dark blue-gray to black fur on the
belly and blackish brown fur on the back. Another type is the cross fox,
which has reddish yellow background fur with a dark brown cross extending
from the nape of the neck to the tail. This form occurs occasionally in
Ciscaucasia and Transcaucasia. Peculiar silver foxes with chocolate-purple
fur are occasionally found in Dagestan. Silver foxes rarely occur in the
Zakataly and Nukha regions in Transcaucasia, only 1-2 specimens being
found in 20,000 animals.
392
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TABLE 79. Geographic variation of skull dimensions (in mm) of foxes from the Caucasian Isthmus*
(396)
Width between ends Number of skulls
Regi d collecti ite Basic length of skull
egions and collection sites asic length of sku of auditory canals ВЕ
Western Ciscaucasia and Don area
> К 131.5 5
ALAS OVIKAWATE AL ии ат <iditetes } 125-138
hi 128.0 5
ро ATCA Neue avers. SN hee twee ole bel He 125-137
(BO7 a. 129.0
а оси НЫ 5
Sal'sk area 193-198
133.5
- i i aes eek = 1
Trans-Kuban Plain, Maikop 128-149 0
Central Ciscaucasia
133.0
feat tas Пос CPoweoa sb Е 3
Kursavka area 124-141 7
Eastern Ciscaucasia
. 134.5
Ipatovo, Beshpagir............- 123-150 7
@rdzhontkidzer\<-e eiew. =) suchen. A 132.0 48
123-141
izl Makh ess eb eaters tees, SMe
Kizlyar, Makhachkala 129.0 FA
120-143
Greater Caucasus
133.0
GVilety ROS We ALO eens. вы. ae eT 4
eee 122-138
Black Sea coast
Tuapse-G ees 6
uaps Е ol 4 blo bo oom . 126-135
Eastern Transcaucasia
121.5
i —__ 14
ОО В nib oo oem aed 115-129
Lesser Caucasus
‘ 125.0
Вог eer ata alte, ie vero eat tee 718-133 3
128.0
Welenovka a copa thoes oer ? 8 7
119-138
Note. Mean values in the numerator, observed ranges in the denominator.
* Material used is in the ZIN and Moscow University collections. Skulls used for measurements were those
of mature males and females, with replaced teeth or teeth in early stages of wearing.
395
Fur traders do not subdivide the fox skins from the western plains of
TranscaucaSsia.into races. These foxes are small and their fur is the
398 usual type of rust in color. As far as the pelts at the Batumi station show,
rust gray foxes predominate in Adzharia; they amount to 71% of all the
foxes. Foxes similar in color and size to the Transcaucasian race number
16%. Foxes of the type transitional from the Yerevan to Transcaucasian
occur on the margins of the Dzhavakhetia Highland (Bogdanovka, Tsalka).
Dimensions of skins and skulls of foxes of some of the races mentioned
are given in Table 78 and 79.
DISTRIBUTION OF
Z L FOX RACES (in %)
h Tbilisi j ; IN AZERBAIDZHAN
Data of 1935-1936
stocks,
Compiled by
N.K. Vereshchagin,
1938
§ Nakhichevan : CO silver fox (4.2%)
` ; E®}sivodushka (4.2%)
\ 7 ] (3 Зстозз fox (2.1%)
|] Е уегеуап
(5) Тгапзсаисазап gray
[1 Тгапзсаисаяап karagan
: ‚ & 0х
% Fur trading stations
FIGURE 164
Analyses of all the data on the distribution of the coloration types and
dimensions of skins and skulls produce a very complex picture of the
geographic variation of foxes on the Caucasian Isthmus. It is clear that
on the Ciscaucasian plains the foxes become consistently smaller and
paler incolor from west to east. In Transcaucasia the altitude factor is
superimposed on this regularity. Nevertheless, the smallest and least
brightly-colored foxes are characteristic of the easternmost populations
inhabiting the hot Kura-Araks Lowland. Another example of the connection
between fur color and landscape are the foxes in eastern Transcaucasia.
396
399
Data on their distribution were obtained from 41 fur stations for the 1935/36
season. The total number of pelts used in the construction of the diagram
(Figure 164) was 28,250. *
From the review of the diagram it is possible to conclude that the
palest-colored foxes occur in the zone of semideserts. Westward, towards
the foothills and mountains, the abundance of bright-colored specimens
increases.
In the Zakataly forest-mountainous region, 4.2% of the furs are silver
foxes, 4.2% sivodushka, and 2.1% cross foxes.
It may be mentioned that to the south and southeast of the Caucasus are
located the areas of distribution of the small races of the same polytypic
group of the Palearctic black-eared foxes: V. vulpes flavescens Gray,
VA vo pies pier se ews ВТ Ve-vulpes Фес op u's’ Blyth:
Taking into account such taxonomic variability of the Caucasian foxes
and following the generally accepted criteria for establishing the area of
origin of the species, the subprovince of the east Mediterranean and
southwest Asia is the area from which the Holarctic fox most probably
originated.
Frequency
MS & BD &®
42 44 46 48 50 52 1 12 13 У 6 7 6 Эш
FIGURE 165. Variation in size of teeth of corsac fox in the Middle Pleistocene
I — length of lower molar row; II — length of Му crown; Ш — length of М›+Мз. 1— Recent
Vulpes corsac, Central Asia; 2 — fossil У. aff. corsac, eastern Transcaucasia, Binagady.
Dots indicate the mean values
The evolution of the skull of the corsac fox was much more pronounced
than in the common fox. The remains from the Binagady asphalt indicate
that the Middle Pleistocene east Caucasian corsac fox was somewhat smaller
than the Recent Ciscaucacian and central Asian varieties (Vereshchagin,
1951b). In addition, the size of Mz and M3 has considerably decreased,
and M3 has even completely disappeared since the Middle Pleistocene
(Figures 165, 166). The normal sized M3 was present in the Binagady
* For the sake of brevity the numbers were not tabulated.
597
corsac fox (43 cases), whereas in the Recent species this tooth is
present in 66% (8 out of 12 cases). The Recent corsac foxes increase
in size to the east, in their presumed country of origin, i.e., the
steppes of the Baikal region and Transbaikalia, where they attain their
maximum size.
Brown bears in the Caucasus have noticeably decreased in size since
the Middle Pleistocene. The Binagady bear from the Middle Pleistocene
beds of the Apsheron Peninsula is closely related to the Recent large
Caucasian bear, Ursus arctos caucasicus, but the Binagady species,
which is probably ancestral to the Caucasian bear, is characterized by
400 bigger molars of a more carnivorous type. This is particularly noticeable
in the shape of the sharp pointed heel of М2. The decrease in the size of
the animals and their teeth towards the Holocene was accompanied by the
flattening of the grinding surface of the teeth with the development of
herbivorous habits. The Binagady Pleistocene bear may actually be
identified as а distinct species (Vereshchagin, 1951с).
The biggest was the Middle Pleistocene bear of the Russian Plain,
known from the Lower Kama (Mysy). This bear, a contemporary of the
Binagady bear, has been identified by us as U. kamiensis N. Ver. sp.
nov. The skull is elongate with a low forehead (Figure 167). In size (basic
skull length 381, 386 mm) it was no
smaller than the cave bears of the
Pleistocene. In the Upper Pleistocene
the bears of the Kama area (Tatar
A.S.S.R.) began to decrease in size
and to develop a convex forehead. The
latter feature is particularly noticeable
in the skulls of the bear
р UA kawkimalkile nsiis) №. Vers sp:
nov. (Figure 167, 2) from the asphalt
VS Boal "ee г © near the village of Nizhnie Kalamalki on
the Shemsha rivulet. In postglacial
time the bears of the Russian Plain
and the Caucasus decreased
considerably in size, as confirmed
by the size of the last molars. Our
diagram (Figure 168), which combines
the elements of the stratigraphic,
geographic and individual variability
FIGURE 166. Lower jaws of corsac foxes in the size of м? of bears, illustrates
1 — Recent, Central Asia, No. 9470, ZIN; these facts.
2 — fossil, eastern Transcaucaasia, Binagady, The Recent populations of bears
No. 23674, ZIN of the Caucasus are genetically highly
heterogeneous. In the present case
it is probably impossible to speak
of the regularities inthe geographic distribution of one monotypic species
since, according to Smirnov's (1916a) studies, the region is inhabited by
the big Caucasian subspecies and two small races — the northern and the
southern, or Mediterranean. The interbreeding of these three forms (the
northern race has probably migrated to the region relatively recently)
makes the studies of the geographic variation of the species very difficult.
398
401
There is а high degree of variability in skulls of badgers from the Middle
Pleistocene of Binagady. Skulls with features of boththe sandbadger (Meles
meles leptorhinus)andthe common badger occur at the same locality.
Most of the badgers, however, were closest to the Recent badger of eastern
Transcaucasia and northernIran—M. meles minor and М. meles
canescens Blanf. It is interesting that the brain of the Recent badgers
has become smaller, which attests to a decrease in metabolic rate
(Figure 169). The first upper molar of the badgers from eastern
Transcaucasia has decreased 8% in width, relative to length, since the
Middle Pleistocene. However, badgers from the Bronze Age of the Armenian
Highland had a wider М! than the early Apsheron forms (Vereshchagin,
1951b).
FIGURE 167. Skulls of bears
1— Recent Ursus arctos caucasicus, Greater Caucasus, No.6169, ZIN;
2—U.karmalkiensis М. Ver. sp. nov, Tatar A.S.S.R., Nizhnie Karamalki, Upper
Pleistocene, No.3, Kazan University; 3—U. kamiensis М. Ver. sp. nov., Tatar
A.S.S.R., Mysy, Middle Pleistocene, No.1, Kazan University
399
The Recent badgers of Ciscaucasia are larger in size, while those from
Transcaucasia are the smallest.
Skulls of Vormela peregusna of eastern Transcaucasia have not
altered appreciably since the Middle Pleistocene. The Recent tiger
polecat decreases somewhat in size from the west to east in the steppe
and desert zones of the U.S.S.R.
The Upper Paleolithic gluttons from western Transcaucasia (Figure 57, 2)
differ from the Recent tundra-taiga species in the deeper mandibles and
larger molars, in this respect exceeding even the older, Middle Pleistocene
forms from the Russian Plain and the Urals (Table 80). This is at variance
with Bergmann's rule, but the phenomenon may be satisfactorily explained
402 by the large size of the local prey: Caucasian ungulates.
Recent Pleistocene
i : 5 6
i
3 +t 2 7
20
и 3
0
Ци ———
30 31 32 33 34 3% 36 37 88 39 40 Я 42 43 mm
FIGURE 168. Stratigraphic and geographic variation in length of М? of
Pleistocene and Recent bears
1—Ursus arctos arctos, Karelia; 2—U.arctos caucasicus,
Greater Caucasus; 3 — 0. arctos meridionalis, Lesser Caucasus,
Talysh; 4—U.arctos arctos (subfoss.), Voronezh, Holocene;
5—U. karmalkiensis М. Ver. sp. nov., TatarA.S.S.R., Nizhnie Karmalki,
Upper Pleistocene; 6— U. kamiensis М. Ver. sp. nov., Tatar A.S.S.R.,
Mysy, Middle Pleistocene; 7—U. arctos binagadensis, eastern
Transcaucasia, Binagady, Dots indicate individual measurements; hatched
circles, mean values
The smallest glutton was collected in the Middle Pleistocene of the
middle Urals.
There are no data on the evolution of the Caucasian martens of genus
Martes inthe Quaternary. However, the Recent martens are represented
in the area by distinct forms. The skull of the Caucasian pine marten
(M. martes lorenzi) is on an average 2.0-2.5 mm longer thar the skull
of the central Russian marten (M. martes ruthena Ogn.); jugal width
is greater in M. martes lorenzi. The Caucasian species is
distinguishable from the Swedish marten (Martes martes martes L. )
by a more highly-developed facial region, a relatively small brain case,
and a narrower postorbital bridge (Kuznetsov, 1941). The coloration of
the fur has more pronounced differences.
400
403
404
These differences should not
be considered significant, even
if it is taken into account that the
species migrated to the Caucasus
relatively recently, as the
differences may be due to specific
features of the Caucasian habitats.
The Recent pine marten varies
little in size and color within its
distribution area in the Caucasus.
Individual and age variability
are much more pronounced, as
expressed, for example, in the
shape and color of the neck spot.
140 100 180 200 220 240) 260mm? This variability, based оп the
material of the Rostov fur station,
is Shown in Figure 170.
The biometric differences
between the Caucasian populations
of the older stone marten
(M. foina nehringi) and the
West European M. foina foina
Erxl. are, as expected, insignificant. Nevertheless, the skulls of these
Caucasian martens are 2-2.5 mm larger than those of the Swiss martens,
and 5-6 mm larger than the skulls of the Crimean martens (М. martes
rosanovi Martino).
The skull of the Caucasian mink (Mtitreola lutreola turovi) is
also considerably larger than the skull of the central Russian species
(Bobrinskii, Kuznetsov and Kuzyakin, 1944). It is known that the increase
in size is fairly characteristic of populations in newly established marginal
sections of an area of distribution.
Detailed morphological and ecological studies are needed for the
clarification of the genetic affinities between the small and big Caucasian
weasels: Mustela nivalis caucasica and M. nivalis dinniki.
These often occur side by side and may therefore be regarded as separate
species.
Cats such as the panther, cheetah, European wildcat and lynx are of
interest from the point of view of their origins and speciation in the
Caucasus.
The size and proportions of the teeth and skeletal elements of leopards
have remained practically the same since the Middle Pleistocene (Table 81;
Figures 61, 6; 85, 1), as have the teeth of cheetahs (Vereshchagin, 1951b).
The Recent Caucasian panthers are smaller, with thinner fur and brighter
coloration in Transcaucasia than in Ciscaucasia (Dinnik, 1914а; Satunin,
1915a). They are larger than the south Asian and African varieties.
The Recent European wildcat of the Caucasus (Felis silvestris
caucasicus) is generally larger than the European variety
(F.s. silvestris): the condylobasal skull length of an adult male of the
former is 99.1 (89.8-102) mm, andof the latter 86-96 mm. The lengths of
the upper tooth row are 33.1 (30-34.7) and 28-32 mm respectively.
Frequency
FIGURE 169. Variation in brain volume of badgers
from eastern Transcaucasia in the Middle Pleistocene
1 — Recent Meles meles minor; 2 — fossil
М. meles aff. minor, eastern Transcaucasia,
Binagady. Dots indicate the mean values
401
(4 03) TABLE 80. Stratigraphic and geographic variation in dimensions (in mm) of teeth and lower jaws of
405
gluttons of the Caucasus and Eastern Europe *
Upper Middle Pleistocene Recent
Pleistocene
oy 2
о ‘bo ees “ > ря -
Measurements es 9 2S of oe a Е Е
2G но ‹ a Е 5 ED за by ь
Е О: о
ЕЕ Meat wks) я = ces «
Е eu. ee a0] Sie us Bani
SRO ao of U a> оо = -о©
Ss u OF Pao нон яя ©
3 > помо < < [2] „4 a оно
оо IN6< HaXx Za elt SCO 2. чо
: 20.5 24.0
Height of jaw behind M,.... о 195-210 710-260
Height of jawnearPm3 .... : 4 17.8 19.3
17.5-18.0 18.0-21.0
1.5 7.0
Теорию рые аи Е. 2) 6 set . é 7 3-1.6. rere
: 4.8 5.3
Width Рот t+ aero te Le ra SIG Е БВ
10.5 130
ПИ Piitguremt-memarets © ie terer : ; . 10.0-11.0 T1.0-11.1
: 6.5 7.4
МЧ ЕТ сое пе о ares Е 0-10. 7028.0
19.5 22.1
Bengtht, М це lene op eae : . . 19.0-20.0 210-232
2 9.0 9.8
Width Ме ayes aie ал 8.0-9.5 “9.0-10.2.
Note. Mean values in the numerator, observed ranges in the denominator.
* Author's unpublished material.
As arule, cats inhabiting the reed thickets of the Lower Kuban are more
uniformly colored than those which live in beech forests. Melanic individuals
occur infrequently in the big beech forests near Nukha, Zakataly and
Madagiz (Lesser Caucasus). In order to make sound conclusions on the
scale of isolation of the Caucasian populations, pelts and skulls from the
north of Asia Minor, the Balkans and the Carpathians must be studied.
The Recent Caucasian lynx (Felis lynx orientalis) is usually
regarded as a subspecies of the European lynx. Its body and skull size
sometimes exceed those of the European form, and its coloration is brighter
than that of the northern form. Morphological changes of the Caucasian
lynx include decrease in size and simplification of the form of the internal
nares in the populations of the Lesser Caucasus.
402
3 Е Га ЕЕ Е Е = 2 :
Е 2 Е + 8 == = ЕЕ Е
7 | Е Е = Е ЕР Е
1 Е 1 Е Е Е == Е
Е НЙ = Е ==: и
2 = = ЕЁ Е he
TAN Sve eee а
22 гея 10 2
\}:
aula a
| wet к yuh,
[ i" \ и ih РР,
| | \ м |
is ‘ ome
i
FIGURE 170. Variation in the size and shape of the throat spot of Caucasian pine martens (1) and stone
cm
[a eS ee ee ee ee ee ee
martens (2). Numbers indicate frequency
TABLE 81. Dimensions of teeth (in mm) of Recent and fossil leopards in the Caucasus
Crown length Му
Locality and age
18.1
25.0
17.8- 19.3
24.5-26.2
Western Caucasus, Recent......... Panthera pardus
Baksan gorge, Sosruko grotto, Early
Holocene (Mesolithic) ..........
P. aff. pardus .
Upper Rion, Kudaro cave, Middle
Pleistocene (Lower Paleolithic).....
P. cf. pardus
Note. Means in the numerator, observed ranges in the denominator.
Order LAGOMORPHA
According to 1. Gromov's measurements (1952), the following
morphological changes have occurred in the European hare of Transcaucasia
since the time of deposition of the Binagady bitumens: narrowing of the
403
406
facial region of the skull, development of more angular orbits, decrease
in the volume of auditory bullae, and narrowing of the ilium. The Recent
European hares of the Caucasus well illustrate Bergmann's rule in that
they increase in size from south to north. The European hares of
Transcaucasia (Lepus europaeus cyrensis) have the following
characteristics: skull length 93-97 mm, liveweight up to 3.5 kg, pale
yellowish brown in color. The skull length of the Ciscaucasian Lepus
europaeus caucasicus varies from 95 to 105 mm, the liveweight
reaches 4.5 kg, and the color is predominantly brownish-yellowish gray
(Bobrinskii, Kuznetsov, Kuzyakin, 1944). According to our measurements,
the condylobasal length of the skull of mature hares (females and males)
from the Kura Lowland in eastern Transcaucasia is 85.3 (79-94) mm (32
specimens), whereas in western Ciscaucasia it is 88.4 (83-96) mm
(25 specimens). The liveweight of mature east Transcaucasian hares
(females and males) is 3.78 kg (14 specimens), whereas the weight of the
Ciscaucasian hares is 4.76 kg (9 specimens). The increase in the size of
the hares is most noticeable in the region of the Khachmas Lowland, on the
Kusary Plain and in the forests of the Lower Samur. This longitudinal type
of variation confirms the southern origin of the species.
Order RODENTIA
Only few species of post-Pliocene Caucasian rodents are known, so that
the phylogenetic lineages of most of them are too short to study their
stratigraphic variation. However, the wide distribution and numerical
abundance of many of the species permits easy tracing of their geographic
variability.
The geographic (latitudinal) variation of morphological and physiological
features of the little suslik is reflected in the increase in size and in the
darkening of color, as well as in the shortening of the period of estivation
from semidesert to steppe habitats.
According to Ognev's measurements (1947), the skulls of the subspecies
Citellus pygmaeus kalabuchovi from the Sal'sk steppes are on
average almost 2 mm larger than the skulls of C. pygmaeus pallidus
from the Kuma region semidesert. The variation with altitude is even more
pronounced: the isolated populations of susliks of Ciscaucasia regularly
increase in size and darken in color with height (Table 82).
From the semideserts on the Lower Terek to the Upper Baksan the length
of the body of the susliks increases on an average by 14 mm, tail length
by 12 mm, foot length by 5 mm, and condylobasal skull length by 3 mm
(Figure 171).
However, the mountain populations of susliks also show considerable
variation in adaptation to hot and dry glacial valleys (altitude 900-1200 m)
and mountain slopes and subalpine meadows (altitude 2500-2800 m). Asa
rule, populations inhabiting the bottoms of the gorges are similar to the
plain dwellers, particularly in color.
The European suslik (Citellus citellus), according to Vinogradov
and Gromov (1952), increases in size and in the relative length of its tail
when the populations are traced from east to west (from the Ukraine to the
Balkans). The Asia Minor subspecies, however, shows pronounced
404
407
morphologic and physiologic variation according to altitude: the body
increases in size and weight, the tail becomes shorter, and the fur color
darker (Table 83). According to Avetisyan (1950), the hibernation of the
suslik in the Alagez area ends by the middle of March, while estivation
begins in the second half of July. In the upland steppes, hibernation ends
only in early April.
TABLE 82. Geographic variation in size (in mm) of little suslik on the Caucasian Isthmus
Lower Terek Dagestan foothills | Terek-Sunzha El'brus slopes
Citellus Citellus Plateau Citellus
Dimensions pygmaeus pygmaeus Citellus pyg- pygmaeus
planicola satunini maeus boehmii musicus
25 specimens 40 specimens 7 specimens 48 specimens
ai ene 206.7 208.5 207.5 220.0
GS АЗ 182-230 182-230 193-215 205-240
ested 30.6 36.3 37.8 les
Taree opie. Lox 30-40 32-43 28-40 45-50
ев 32.1 32.1 33.2 37.4
a mene оне La 21-35 30-36 28-34 36-38
Gonduiepasdlicleftlidencth Deed eee pol, «ЕР OO
Lah inal, hte 39.5-42.0 37.9-42.6 37.8-41.9 425-452
Note. Means in numerator, observed ranges in denominator.
Some general evolutionary trends and specialization rates can be studied
in the jerboas (genus Allactaga).
TABLE 83. Changes in size (mm) and weight (g) of Asia Minor suslik (males and females) at different
altitudes
Zone 1550-2190 т
above sea level
Zone 1255-1550 m
above sea level
Dimension
203 215
Вора ое Е 9 р МЕНЕ pee
175-230 180-390
а ель вами yee ftaeeote ae ae лы al ES a
33-37 21-59
МЕРЕ, Rt fll. а 223% 4 НЕ TSM
188-372 184-430
Note. Means in the numerator, observed ranges in the denominator.
405
408
42 44 46 48 50
Astrakhan
Manyoh
© Stepnoi
oO
я
Pp
5
„^^
я
L
on)
my
Budennovsk RA
Terekli-M aS
~Stavropol
у
(
Ce
Kizlyar 9
Pyatigorsk
и® Sulak
\Mik -
ye ew, Grozny © Khasavyurt @
©) \
Mt. ЕГЬ
at Ee
Г Derbent ©
тео ое т pate
FIGURE 171. Variation in size of little suslik in Ciscaucasia. Rectangles indicate the length of a foot
squared. Populations from the Lower Terek taken as an entity. Black dots — points of trapping
Makhachkala
(©)
The jumping ability of Recent jerboas of Ciscaucasia and the Apsheron
Peninsulahas improved since the Middle Pleistocene (Binagady) due to the
increased length of the tibia relative to the femur, as shown in the increase
in the jumping index (length of tibia/length of femur X 100) of the small and
Asia Minor jerboas (Table 84).
I. Gromov (1952) has pointed out that the structure of the metapodia in
the great jerboa from the Binagady asphalt (Allactaga jaculus
bogatschevi) is more primitive than in the Recent form in the wider
spacing of the distal part of the foot and in the longer free end of the second
metatarsal (11442).
Hamsters (Cricetinae) have occurred in the Caucasus since the Middle
Miocene. It is difficult, however, to establish the affinity of the Miocene
(Palaeocricetus, Belomechetka) and Pliocene (Stavropol) hamsters
with the Quaternary genera Cricetus and Mesocricetus. Hamsters
from the Lower Pliocene alluvium of Stavropol (Kosyakin quarry) are
characterized by large molars. The hamster from the Binagady asphalt
(Mesocricetus auratus planicola) is intermediate in size between
M. auratus raddei and M. auratus nigriculus. The Binagady
species differs from the forms of Mesocricetus known inthe U.S.S.R.
in its hard palate and short incisor foramina. The structure of the
postcranial skeleton of the Upper Pleistocene hamsters from the Chiaturi
manganese region indicates that the population was no less specialized than
the Binagady population.
406
TABLE 84. Changes in jumping index of Middle Pleistocene jerboas*
Jumping index
Species
Binagady Recent
дНтасваеа jaye lise. pater el ве 125.0 127.0
IS У Е Зоо ие ро Oy) о о о 126.1 129.5
ево ба обовоо в ово 129.5 136.8
* Ме calculated the indexes from the mean values given by I. Gromoy (1952); he tried to reconstruct
the ancient landscapes of the Apsheron on the basis of certain morphological features of the Binagady
jerboas. His assumptions, however, on the existence of either soft ground, mosaic landscapes or
diversified relief at Binagady are poorly founded.
The geographic variation in the Recent populations of hamsters in Asia
Minor is mainly reflected in changes in the depth and brightness of the
rusty color of the back and sides, and the black color on the abdomen
(Argiropulo, 1937). Also the mountain populations are, as arule, larger
than those of the Ciscaucasian foothills, though the ecological variability
of the species is great. On the high Armenian plateaus, hamsters begin
to hibernate at the end of October or early November, whereas on the lower
Kakhetian plateau they begin only in early December.
The Recent gray hamsters of the Apsheron Peninsula have developed
a narrower anterior palatal region, larger molars, a more protruding
coronoid process, anda Slightly recurved lower incisor and Mg with
shortened heel. These changes have occurred since the Middle Pleistocene.
The functional significance of these minor changes and the evolutionary
trends of the species are difficult to explain. It is simpler to record the
facts, as has been done by I. Gromov (1952).
The process of morphological and ecological evolution of the species as
polymorphic as the house mouse (Mus musculus) in the Caucasus is
409 very complex. This species was widely distributed in the Middle Pleistocene
of eastern Transcaucasia (Binagady), and also occurs in this region today.
The morphological and ecological differentiation of the species has led to
evolution of the following subspecies: white abdomen and short-tailed
(steppe and semidesert forms), M. musculus hortulanus,
M. musculus tataricus; gray abdomen and long-tailed (mountain-
forest forms), M. musculus abbotti, M. musculus formosovi;
and introduced domestic forms, M. musculus musculus. This
variability is as developed as the variability of the common field mice of
the Caucasus (discussed below); however, it cannot yet be studied using
the modern methods of morphological investigation (see Geptner, 1930;
Gulii, 1930; Sviridenko, 1935а).
The systematics of the common field mice of the Caucasus (genus
Apodemus) has not been extensively studied, but on the basis of the
studies of Argiropulo (1940a, 1946), Sviridenko (1936), Kuznetsov
(Bobrinskii, Kuznetsov and Kuzyakin, 1944), Shidlovskii (1953a) and our
observations, the genus may be considered as being subdivided into four
407
410
species: А. mystacinus, A. cflavicollisy,An-fulvipectws tand
A. sylvaticus (see also Chapter V). This classification is tentative
and subject to future experimental ecologic studies.
The Asia Minor mouse (A. mystanicus), distributed in the Balkans,
Asia Minor and western Transcaucasia, is considerably smaller in size
in the Caucasus than in the Balkans. According to Shidlovskii's
measurements (1953a), the condylobasal length of the skull of the Balkan
specimens is 28.4 mm (34 specimens), and of the Transcaucasian specimens
26.5 mm (200 specimens). The length of the upper molar row is 4.9 and
4.5 mm respectively. The Balkan populations are lighter in color than
those in Asia Minor and Transcaucasia. According to Shidlovskii (1953a),
this animal migrated to the southern slopes of the Greater Caucasus
(Map 55) recently, during the last few decades. In the present case, the
body size of the animal, which migrated into a new mesophytic environment,
has decreased considerably. Paleontological studies of the Imeretian caves
will clarify whether this is a primary or a secondary immigration and will
consequently shed light on the degree of morphological divergence of this
east Mediterranean species.
The picture of the geographic variation of the other three species on the
Caucasus Isthmus is much more complex. The yellow-necked mouse of the
European type, with feet 24-25 mm long (close to A. flavicollis
Samariensis) occurs only in the river valley forests on the Lower Sulak
in eastern Ciscaucasia. * In the far southwest the southern boundary of the
distribution of this species reaches the Lower Don (Map 55).
The relatively small A. fulvipectus**, with a foot 22-23 mm long,
occurs in the deciduous forests of the lower mountain belt in the Krasnodar
area, in Abkhazia, Adzharia and northwestern Azerbaidzhan, and insome places
in northern Armenia (Map 56). This species is easily distinguishable from
the common field mouse, even in juvenile states, by its larger dimensions,
greater length (relatively shortened) of the foot and the presence of an oval
yellow spot on the breast. The species comes out into the sun to feed only
rarely. It is a good climber and often lives in hollows in trees. It also
differs from the common field mouse in its weaker response to light.
It seems likely that the relatively short foot and tibia, the longer tail of
the yellow-spotted mouse and its relatively narrow internal nares
(Argiropulo, 1946), as compared with these features of the common field
mouse, are adaptations to slower movement and better climbing.
The populations of this species vary even within their area of distribution.
As arule, the southern forms are somewhat smaller, while to the east
their size increases. Morgilevskaya (1954) has published data on five mixed
populations of the yellow-spotted and common field mice. According to
her data, the condylobasal length of the skull is (accordingly to locality):
Sukhumi — 23.8, Gagry — 22.3, Batumi — 23.1, Akhalkalaki — 22.7,
Lagodekhi — 23.7 mm; length of hind foot: Sukhumi — 22.6; Gagry — 21.3,
Batumi — 21.4, Lagodekhi — 22.7 mm.
” This is true only if the labels on the material in the Moscow Zoological Museum have not been mixed.
Geptner and Formosov (1941) have identified 5 specimens from this region and I studied one specimen,
but the origin of this species is not yet clear to me.
“* We have assigned to this species the Caucasian subspecies А. flavicollis ponticus, A.flavicollis
parvus and A. sylvaticus fulvipectus of earlier authors.
408
411
Populations of yellow-spotted
mice are common in a number of
valleys in Central Ciscaucasia
(Urukh, Ardon, Terek) and on the
piedmont plains of eastern
Ciscaucasia. In this region the
animals vary considerably in size
from one valley to another, which
is due to both genetic isolation and
differences in habitats. The higher
degree of ionization of the air in
the area of tectonic fracture has
been suggested as the reason for
the larger size of the mice in the
Armkha ravine (tributary of the
Terek) (Ushatinskaya-Dekalenko,
1933). If this is true, then the size
of mice in the mountain populations
may be an indicator in surveys of
radio-isotopes.
In eastern Transcaucasia this
Species primarily occurs in the
piedmont lowlands. Careful
measurements in а series of
preserved specimens of
A. fulvipectus and common
field mice of the same age group
gave the data on the geographic
variation in foot length, as shown
18 19 20 af 22 23 24 25 26mm in Figure 172.
FIGURE 172. Variation in foot length of common field Isolated bones of the common
mice on the Caucasian Isthmus field mouse (A. sylvaticus
Apodemus flavicollis; 1— Rostov region. subsp. ) occur in the Middle
А. fulvipectus: 2— Caucasian Reservation; Pleistocene asphalts of the Apsheron
3 — Alazan-Agrichai valley (Transcaucasia); Peninsula. No morphological
4 — Lesser Caucasus (Delizhan); 5 — Talysh. differences between these and the
A.sylvaticus: 6 — Rostov region; 7 — Caucasian Recent forms have been recorded
ОН 8 я В Ossetia (Lars, fam ane in this material. In Ciscaucasia,
— Alazan-Agrichai valley (Zakataly ), 10 — ee annie species те widely АЕ
Araks valley. On the left — foot of A. sylvaticus; : ; é
on the right — А. fulvipectus. Ordinate — on the plains, in the foothills and
number of specimens in the mountains, particularly in
areas of sparse vegetation (Map 57).
The mountain populations of this species are characterized by higher counts
of erythrocytes in their blood (Kalabukhov, 1940). In Transcaucasia the
common field mouse is somewhat larger, which sometimes makes it
difficult to distinguish from A. fulvipectus. A yellow spot is almost
always present on the breast of mature specimens.
The Caucasus is thus an ancient and self-contained center of speciation
of domestic and field mice; this confirms their Pliocene origin in this
country.
409
412
The red-tailed gerbil from the Binagady bitumens (Meriones
erythrourus intermedius) is a form intermediate between the Recent
gerbils occurring to the west and east of the Caspian.
The primitive features of the Middle Pleistocene gerbil are reflected
in the shortness of the tibia, which is only 113.8% the length of the femur,
as compared with 125.9% in the Recent gerbils of the Apsheron. *
Insofar as the lengthening of the tibia relative to the femur indicates
an adaptation for jumping (jerboas), it can be stated that the Recent gerbils
became ''jerboaized'' (N. A. Smirnov's term) since the Middle Pleistocene.**
Evolution in space and time can be studied inthe case of the Caucasian
voles (genera Prometheomys, Arvicola, Microtus) as rewardingly
as in mice.
The teeth and lower jaws of Prometheomys from the Acheulean beds
of KudaroI and the Upper Paleolithic beds of the Gvardzhilas cave are not
noticeably different from the Recent forms. This is understandable, since
the morphological features of burrowing rodents reflect conservativeness
in habit. The data on the geographic variation in the color of Recent
populations are summarized in Table 85.
TABLE 85. Variation in abundance of melanic varieties in the distribution area of Prometheomys
Upper Laba and Belaya Upper Bzyb Upper Terek
Grayish brown.......
* Collections of ZIN AN SSSR.
The increased proportion of the melanic varieties is probably restricted
to the western part of the distribution range.
The Recent populations of water vole (Arvicola terrestris) vary
noticeably within the Caucasian and adjacent regions. At least 9 varieties
have been described in the Caucasus, and not less than 6 varieties from
neighboring areas in Iran, Turkey and the Russian Plain. It is difficult,
however, to trace the geographic variation in the morphological characters
of the species.
As arule the northern marshland forms from the Lower Don, Kuban
and Volga are somewhat larger than the mountain and south Caucasian
forms (Table 86).
Populations of this species from some sections of the distribution range
are characterized by minor morphological adaptive features. The upper
incisors of the mountain ''burrowing'' populations of Balkaria, Ossetia,
and Dagestan resemble those of mole voles. The curvature of the incisors
is greater than in the marshland forms, and the degree of differentiation
* Indices of the mean values were computed by us from the measurements given by I. Gromov (1952).
** 1.Gromov has also indicated that the Binagady gerbil was of a “more running" type (i.e., less of a jumping
type) than the Recent species.
410
of the fur is less marked. The change in shape of the incisors is due to
their adaptation for digging.
TABLE 86, Variation in skull size of the water vole in the Caucasian Isthmus*
Lower Don Lower Kuban Transcaucasia
Eastern Dagestan
Arvicola Arvicola Arvicola Arvicola
Dimensions terrestris terrestris terrestris terrestris
tanaiticus cubanensis kuruschi persicus
10 specimens 11 specimens 32 specimens 32 specimens
41,2 41.0 38.5 39.6
Condylobasal length of skull... 41.0-41.5 39.5-42.6 38.2-38.3 37.3- 41.9
10.2 9.9 10.1
rs —_—_ 4 —_————_
Length of upper molar row 10-10.4 9.2-10.5 9 9.6-10.8
Note. Means in the numerator, observed ranges in the denominator.
* Measurements by Орпеу (1933).
Up to 8 subspecies of the snow vole known from Palestine to Kopet-Dagh
are also found in southwest Asia, the Caucasus and Central Asia, The
geographic variation of the Asia Minor snow vole occurring between the
Lesser Caucasus and Kopet-Daghis expressed in lighter coloration correlated
with increasing draught and isolation, in the shortening of МЗ along the
west—east gradient, and in a general decrease in size (Table 87).
(413)
TABLE 87. Geographic variation in size (in mm) of Asia Minor snow vole*
nt Oe 2 5 ai ke eaten
Dimensions 5 a 2.42) 9c o м ой вв о 2
ЗЕЕ! ВР еяЕ Soap a Eee 2
OVE Eo Se | Cea iene SOLON Ne ns OB Bice
сена
Be sf mwea elle ok | eek eB еее
м Е а Ее мЕБЕЗО
р 109.4 QLD 108.2
Body length......... 99-121 109-142 97-120
й 58.1 54.0 59.3
а sn Gio elles iol 59-68 32-90 51-60
ль 18.8 17.9 aoe
oot length........ . 12.2-20.0 13-20 17-20
26.4 21.8 211.8)
Total skull length 5 .0-27.8 26 3-29.6 25 .8- 28.5
+ 1 5.9 6.3 RACE IT,
Length of upper molar row 55-62 6.1-6.7 5.5-6.5
Note. Means in the numerator, observed ranges in the denominator.
* Author's measurements on the ZIN material.
411
413
414
The snow vole from the Greater Caucasus is the smallest variety,
indicating that humid mesophytic regions are not suitable for this animal.
The decrease in size of this species towards the southeast, and its
disappearance from the intermediate heights of the southern slopes of the
Armenian uplands, indicate that the southeastern sections of the present
distribution area are in a state of decline. This also indicates that in the
past a more humid phase existed, during which the distribution area of the
species extended further to the east.
The geographic variation in the Caucasian snow vole on the Greater
Caucasus is reflected in the decrease in size and lighter coloration from
west to east. The change in color is probably due to the fur becoming
bleached in sunny woodless Dagestan. The inversion of the zones in the
central part of the Greater Caucasus is somewhat superimposed on the trend
of brightening of the color.
The decrease in size from west to east is shown by the measurements
given in Table 88.
It should, however, be mentioned that size variation of this species of
vole has not yet been traced through the altitudinal and landscape zones.
TABLE 88. Geographic variation in size (in mm) of Caucasian snow vole”
Eastern Caucasus,
Western Caucasus Central Caucasus
р я Dagestan,
d f Microtus gud Microtus gud г
Dimensions ‘ P Microtus gud
nenjukovi gud :
lghesicus
128 specimens 116 specimens
8 specimens
Body lengths. еее, 120-152 102-149 100-121
Taildenptlijewn-ee cet) Ae 68-106 61.0-85.0 58-78
о ПОЙ поме ос овес 19.0-26.9 18.0-23.0 18.5-21.2
Condylobasal length of
TE SS oe соо 26.4-28.6 25.0-29.2 26.4-27.2
Length of upper molar
SOS meee АЕ 6.7-7.8 6.0-7.0 6.0-6.6
* Measurements of Орпеу (1950) and author's (for Dagestan).
The Middle Pleistocene Apsheron vole (Microtus (Pitymys)
apscheronicus) differs from the Recent pine vole of the Caucasus in its
larger size, its longer tooth row (6.3 mm as compared with 5.7 mm of the
Recent species) and in details of the shape of the lower jaw. It is also
possible that the extinct species was of a less mesophilous character than
the Recent form.
Geographic variation of the Recent pine vole is poorly developed. The
subspecies described for the Caucasus reflect the subjective approaches
of investigators or the ecological variants (and age groups) arising due to
variation in living conditions from year to year. The subspecies described
are: M. majori majori from northeast Asia Minor, the Kars Highland,
Dzhavakhetia, Mingrelia, and the central and western parts of the Greater
Caucasus; M. majori ciscaucasicus from the northern slope of the
1704 412
415
Bol'shoi Range, inits мез{егп ап4 сепёга]рагёз; М. majori suramensis
from the central sections of both slopes of the Bol'shoi Range, Surami
and the Lesser Caucasus; M. majori dagestanicus from inner
Dagestan, Tushetia, northwestern Azerbaidzhan, the northeastern slopes
of the Lesser Caucasus and Dhavakhetia; M. majori schelkovnikovi
from the forests of Talysh.
TABLE 89. Geographic variation in size (in mm) of common vole in the Caucasus"
n n nn n
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Е Е а a а += fey TAY с) е Я эх
Di , Е зна м Е ма BS sy
imensions a ® 22 8 в = оо ra 2)
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ное = of on & MOMs а ооо
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№ © — ох Gow t fac HSE
Condylobasal length of 26.3 23 .8-27.3 25.3 25.2
УЧИТ Go sab asso6000 25 .8-27.3 j ; 24.0-26.9 24.0-26.0
Length of upper molar 6.0 5.7 5.9 5.5
WOW Боро вон oe oe 5.8-6.5 5.3-6.2 5.2-6.0 5.2=5.7
17-18.2 15.5-19.6 14.1-18.0 0.
Foot length.......... -18. 0-19. 1-18, 15,0-17.0
Note. Means shown in the numerator, observed ranges in the denominator.
* Measurements of Krasovskii (1930), Ognev (1950) and author's (for Talysh ).
The Talysh specimens are distinguishable by their intense rusty color
on the sides and their small size. М. majori dorothea Ellerm. from
the Elburz Range is probably synonymous with the latter subspecies. The
pronounced differences between the M. schelkovnikovi subspecies
and the pine voles of the Lesser and Greater Caucasus attest to the long
period of isolation of this animal in the southeastern section of its
distribution range. The overlap of the distribution ranges of the former
four subspecies is self-explanatory. The rather vague systematic diagnosis
of the forms described indicates that in order to understand their geographic
variation special ecologic and morphologic studies must be made.
It is known that the common vole (Microtus arvalis) decreases in
size from north to south on the Russian Plain. On the Caucasian Isthmus,
with its diversified relief and inversion of the zones, Bergmann's rule does
not strictly apply to the distribution of the species. Studies of the geographic
variation of this species must be done systematically across the altitudinal
and landscape zones, taking into account seasonal and annual variation.
Some numerical results of the studies of the Caucasian subspecies are
given in Table 89.
Animals inhabiting the dry longitudinal valleys east of the El'brus and
inner Dagestan are characterized bya lighter color than those inhabiting
the humid foothills.
413
The type of geographic variation described and the characteristic features
of the area of distribution attest to the ancient origin of the species on the
mesophytic uplands of the Mediterranean.
The Middle Pleistocene steppe vole, Microtus socialis, from the
Binagady asphalt is almost identical in skull structure to the Recent species
(Gromov, 1952).
According to Ognev (1950, p. 400) these subspecies increase in size from
north to south; their auditory bullae also increase in size and the structure
of M and МЗ becomes more complex. However, these facts need further
study, particularly in relation to the specific ecologic features and to height
above sea level. Considering the great size of the total area of distribution,
the variation mentioned is not particularly significant.
All this indicates that the relationships between the steppe vole and the
xerophytic landscapes of the Mediterranean have remained stable; they
date at least from the Lower Pleistocene.
Order PROBOSCIDEA
In the south of the Russian Plain, in the Caucasus and in southwest Asia
(as in the Mediterranean in general), the first known true elephants are
represented by highly specialized forms: Elephas (Archidiskodon)
planifrons, E. (Hesperoloxodon) antiquus, E. (Archidiskodon)
meridionalis from the Middle Pliocene (Trouessart, 1898-1899c;
Pavlova, 1910a; Bogachev, 1923-1924; Osborn, 1942; Simpson, 1945).
The smallelephant (Phanagoroloxodon mammontoides), as
described by Garutt (1957a) from some Upper Tertiary beds of western
Ciscaucasia*, is probably of the same age. The affinities of this species
to those mentioned above are not clear. However, the species is considered
to be close to E. (Hesperoloxodon) antiquus or E. (Parelephas)
trogontherii, so that Garutt's identification of this form as Early
Pleistocene is undoubtedly false. The absence of direct ancestors of
the species listed in the Oligocene, Miocene and Lower Pliocene beds
is probably not due simply to change. It is worth noting that the
remains of elephants definitely do not occur in the immediately post-Pontian
beds in Ciscaucasia and in the northern Black Sea area as they do, in the
alluvium of the Kosyakin quarry near Stavropol and in the clay fissure-
fillings in the Pontian limestone, exposed in the Odessa catacombs. In spite
of the ''taphonomic universality'' of both burials, the proboscideans are
represented only by mastodons (and Dinotherium at the Kosyakin
quarry).
Evolution of the elephants in the region must be studied mainly from
tooth material, on which the poorly-developed hypothesis of the evolutionary
lineage .E. planifrons-E, meridionalis Е. trogontuneritt=
416 mammoth has been based. The systematic position of the so-called ancient
* The Phanagorian elephant has been described fiom a permineralized skull (with broken teeth) of °
unknown locality in the Krasnodar museum. Garutt's (1957a) photographs show teeth of elephants
collected from the conglomerates nearSennayaand Akhtanizovskaya on the Taman Peninsula, but his
identification of the material as belonging to the Phanagorian elephant is not certain; these teeth seem
rather to belong to the southern and ancient elephants (Vereshchagin, 1957а). There is unfortunately
no space for discussion of the opinions of Sherstyukov (1954), which are lacking in clarity.
414
417
elephant [Е. antiquus] is not sufficiently clear. Russian paleontologists
have either identified this form with the Loxodonta group or have related
itto E. planifrons and E. meridionalis. It has been identified
as Palaeoloxodon by Dubrovo (1957). Because of the scarcity of
remains of the ancient elephant, it is only possible to construct a
distribution map (Map 76). Over 400 casts and descriptions of various teeth
of Upper Pliocene and Quaternary elephants, collected from the Russian
Plain and the Caucasus, have been studied by us in the Russian museums
and collections. In addition to distribution maps showing occurrence of
teeth (Maps 74-76), we can give a general description of the teeth of various
phylogenetic grades and a summary in table form of the main morphological
features. Most teeth and bones of Pliocene and Quaternary elephants in the
Caucasus occur in redeposited sediments. All the material has been washed
out of the primary bone-bearing lenses by streams and redistributed in
conglomerates, gravels and sands. The localities with Е. planifrons
and E. meridionalis in Ciscaucasia are situated in delta zones of
ancient streams and mudflows, which once flowed from the Greater
Caucasus to the Akchagyl and Apsheron seacoasts. The bones often occur
in ferruginous gravels and sands of the high (third or fourth) river terraces
on the inclined piedmont plains. According to Bogachev (1923-1924), most
of the bones of southern elephants on the coast of the Sea of Azov (Lower
Don) have been washed out of the Middle Pliocene beds and redeposited in
Late Pliocene gravels.
In eastern Transcaucasia the bones occur mostly in loesses, loams
and gravels, around the ancient coasts of the Kura bay of the Akchagyl and
Apsheron seas. Оп the Lesser Caucasus uplands, bones of Upper Pliocene
and Lower Quaternary elephants occur in river and lake-river sediments,
like those near Leninakan and Erzurum. Elephant bones occur at Tsalka
in conglomerates between doleritic lava flows, dated as Gunz and Gtinz-
Mindel (see also Burchak-Abramovich, 1951а).
Teeth of, Elephas planifrons* and Ев, meridionalis are
relatively small, with short and weak roots. Their enamel pockets are
coarse-ribbed, comb-like plates, formed of enamel cylinders intergrown
longitudinally. With a covering of dentine and cement, these widely-spaced
pockets form enamel bellows; upon wearing, the pockets became
transversally flattened, forming enamel loops with thick walls protruding
5-6 mm above the surface in mature individuals (Figure 173). The working
surface of the tooth was used for grinding large pieces of plant material.
The bases of the enamel pockets, near the pulp cavity, form a uniform
roller with poorly-developed papilla. The cement of the teeth of these
early elephants has been altered and weakened to such an extent that the
On the Russian Plain remains of Upper Pliocene elephants occur mostly in
the south, none having been recorded north of the latitude of Moscow and
Kazan.
As far as diet is concerned, E. planifrons and E. meridionalis
were probably still fairly close to the mastodon. Striations and scratches
in the enamel of the teeth of these elephants strongly resemble those on
mastodon teeth, though in mastodons the grooves are usually transverse
* Саги (1957b) has expressed an opinion, not yet well founded, that the so-called Е. planifrons
known from the U.S.S.R. is merely an early form of Е, meridionalis,
** The cement and dentine of these teeth usually resemble alabaster and marl.
415
to the jaw, while in elephants they are longitudinal. This is probably due
to differences in the mode of grazing. The grooves were undoubtedly formed
by sand grains and small stones taken up together with roots and grasses.
The seasonal feeding of elephants in the thickets and swamps and along
the shores of lakes and in river valleys undoubtedly explains their relatively
frequent burials in water-laid sediments.
Tooth replacement in Upper Pleistocene elephants has not yet been studied
thoroughly, but it is generally assumed that in each jaw 5 molars were
worn off and replaced by the appearance of new molars from behind and
above in each jaw, in a manner similar to the replacement of the molars
of the Recent Indian and African elephants. The last, 6th tooth, the biggest,
with thickest enamel, was characterized by the absence of a posterior
crushing surface; it functioned only in the last stages of the animal's life.
This type of molar replacement, which was probably also characteristic
of mammoths, is taken into account in the discussion which follows. *
Towards the Lower Anthropogene, the number and size of enamel pockets
and consequently the height of the teeth have considerably increased; the
enamel wall has become thinner and less folded. The teeth, particularly
the lower teeth, have become relatively narrower and longer, and their
volume and weight have increased both absolutely and relatively. This stage
in the morphogenesis of elephants is represented by a series of burials on
the lower (second and third) river terraces of Elephas (Parelephas)
wusti and Е. (Parelephas) trogontherii.** The occurrences of
bones of E. trogontherii are best known in the gravels of the third
418 terrace on the left bank of the Dniester, near Tiraspol, and in the Araks
valley, near Yerevan and Leninakan (Map 75).
0 5 10 ст
FIGURE 173. Section through М5 of the southern elephant from the northern coast
of the Sea of Azov
Section shows enamel pockets and specific features of loop formation (a — loop;
b — pocket)
Fragment of the skull of a southerm elephant from Khapry (collections of Institute of Geological Sciences
AN SSSR) shows that the replacement of the second upper tooth by the third took place directly from
above, similarly to the mode of replacement in mastodons, rather from than above and behind as in
mammoths and Recent elephants.
The latter name poorly reflects the actual situation, since the increase in abundance and area of
distribution of the "trogontherial"” beaver occurred probably as early as the Upper Pliocene.
mm
416
419
The Middle Pleistocene sees the beginning of the final stage in the
evolution of elephants. It coincided with the rapid development and
proliferation of Middle Paleolithic man, who rapidly perfected the art of
hunting, includingthe elephant hunt. By that time E. trogontherii had
decreased in size and evolved into the mammoth, E. (Mammonteus)
primigenius, and had completely changed in appearance, having
developed a woolly coat*, longer and more sharply curved tusks, and
stouter metacarpals and metatarsals. The evolutionary trends in the teeth
included closer spacing of the enamel loops and increase in their
number per unit length, thinning of the enamel, and smoothing of
the enamel surface (Figure 174). Sharp edges developed on the bottom
of the enamel pockets; pointed papillae situated along the sharp rim
protruded into the pulp cavity. The teeth roots developed into long,
curved or straight hollow tubes and the closely spaced enamel
lamellae make a finer, and possibly more durable, grinder. Traces of
vegetable matter are found only rarely on the truncated surfaces of the thin
enamel of the teeth of mammoths and E. trogontherii. While feeding
on twigs and tree branches undoubtedly continued, grass as a diet began
to gain in importance, as has been definitely established by the contents of
the stomachs of the frozen mammoths in Siberia.
Remains of true mammoths inthe Caucasian Isthmus are more common
in Ciscaucasia than in Transcaucasia. As as rule, they occur in the young
(first and second) river terraces. The state of preservation of the enamel,
dentine and cement of the mammoth teeth is always strikingly different
from the preservation of these components in the teeth of earlier elephants,
as roots andthe pulpcavities of mammoth teeth are often preserved in their
entirety.
The earliest remains of mammoth-type elephants from the Caucasus
were found in the Girei quarry on the second terrace of the Kuban. Only
the materials from the П'зкауа site in the Trans-Kuban region have been
dated. Four fragments of molars, many pieces of tusk, and up to 40
assorted incomplete skeletal elements including tarsals and carpals have
been collected from this site.
The last occurrence, stratigraphically speaking, is part of a left upper
jaw with a very worn 6th tooth from the travertines of the northern slope
of Mt. Mashuk. The whitish enamel, the light cement—light brown
color of the bone in fresh fracture, and the characteristic smell of raw
bones upon scratching — all these features are highly unusual for a burial
so far south. The small size of the tooth, the thin enamel (1.0-1.2 mm), and
the small diameter of the tusk indicate that the fragment belongs to a
stunted, very late variety of mammoth of Late Paleolithic and even Neolithic
age.
In Transcaucasia, mammoth teeth have so far been found only in the Gori
depression (Burchak- Abramovich, 1946; Gabuniya, 1952a). The tooth
described by Gabuniya (1952a), an unworn М*, was collected on the first
terrace of the Kura River in the town of Gori. On the basis of the small
size of the tooth with 12 lamellae over 10 cm and enamel thickness of 1.5mm
Gabuniya has related the tooth to the ''stunted'' mammoths of the west
Mediterranean. On the basis of the mode of its occurrence the specimen
has been dated as early post-Wurm, i.e., Lower Mesolithic to Neolithic.
* So far there is no information on the woolly coat of E. trogontherii.
417
Gromova (1948) has assumed, without any evidence, that ''some mammoths
lived in the Caucasus in pre-Riss time.'' Burchak-Abramovich (1946)
considered (incorrectly) that the Transcaucasian mammoth is a
representative of the northern tundra. At variance with these investigators,
Gabuniya has correctly emphasized the probability of evolution of small
local forms of mammoth in the south of eastern Europe and in Transcaucasia.
FIGURE 174. Longitudinal sections
1- м8; 2 — МвоЁ mammoths from the Russian Plain
418
Even if the northern mammoths migrated to the Caucasus during the
maximum height of the cold periods, it is doubtful that they could migrate
farther south than Ciscaucasia. In the Upper Pleistocene the western
migratory route of the northern mammoths through forests and steep rocks
was difficult, and the eastern route passed through a country too dry and
hot for mammoths.
420 Thus Transcaucasia was probably the southern boundary of the
distribution cf the mammoths, and this is confirmed by the absence of
reliably identified remains of mammoths from the Armenian Highlands,
Iran and Southwest Asia in general. The xeromorphic environment with
poorly-developed wood vegetation, which evolved (at least in the low valleys
of these regions) towards Middle Pleistocene time, was not suitable for
the elephants of late and northern types.
North of the Ciscaucasian plain, mammoth remains are the most common
paleontological finds in the valleys of the Don, the Volga and the Ural, and
farther north the proportion of mammoth bones also steadily increases by
comparison with the bones of earlier elephants. The results of our counts,
done in 1950-1955, are given in Table 90.
TABLE 90. Number of teeth of fossil elephants in regional museums
Mammoths
Elephas
trogontherii
Elephas
meridionalis
ANNI 5 og ob оъяноюовов
ОЕ а оо оо ое о в 33*
Volo gda |. о асы 23*
Seven) ОИ ИЗЕЗИАИ ооо Goo b a oy Ob 59
(ОЕ 9 о ооо бор оо ово о 66 10
ОЗУ Бо alo бо ооо 9яаосо
SEMAN, dale blak 6 5 За чо
Temryuk and collections from
Taman and at the Paleontological
OGRE (о о о о ое моб hob OSG .
ЧУ 6 бр ово юобосбоворов
SUNOCO 6 Зо бб бо соборе еее с
INOS G45 6 al ао ово Зов
ЧЕ ЕЕ оны боб обо лов
Note. Asterisk indicates cases when bones other than teeth were counted.
The distribution of the species, as given in Table 90, indicates more
than the apparent decrease in the frequency of burial of mammoths in the
south and the increase in erosion and exposure of older beds containing
remains of southern elephants. Recorded distribution of the proportions
of the species is due to the fact that the mammoths were very abundant in
the north, the main areaoftheir distribution, whereas the distribution areas
of E. trogontherii and E. meridionalis were further south.
419
(421)
FIGURE 175. Print of worn surface (enamel loo eal nd outlines of teeth M of fossil elephants
of the Russian Plain and the Caucasus (M°— a; b)
Ele ss (Mammonteus ) ae a la, 1b — Sukhona valle iv No. 10401, pete
Aas nae peat in Don valley, un-numbered, Vor = Mu За — Kir oo ee
Mode aleolithic site near Kiev, No.302, eh of Zoology o ee the me ‘ie env f Sci
of th nn 5. 5$. К
420
(422)
FIGURE 175 (continued )
4b — Middle Paleolithic site, Chokurcha, in the Crimea, un-numbered, Simferopol Museum;
da — second terrace of Kuban, Girei quarry, near Kavkazskaya, un-numbered, Armavir
Museum; ба — Terek valley near Naurskaya, un-numbered, Georgian Museum; 7a, 7b — Khazar
age of Volga valley, Chernye Yar, un-numbered, PIN [Paleontological Museum] ("Гате”
Elephas trogontherii and "Early" mammoth);
421
(423)
FIGURE 175 (сопипиеа)
Elephas (Parelephas) trogontherii; 8a,8b— third terrace of Dniester at Tiraspol,
un-numbered, Moscow Geological Prospecting Institute. Elephas (Archidiskodon)
meridionalis; 9a — Araks valley, Leninakan, Kazachii Post, un-numbered, Geological
Institute AN Arm.S.S.R.; 10a,10b — conglomerates of Taman Peninsula, No. 1249/222,
№. 1249/42, PIN (Phanagorian ('? ) elephant);
422
(424
‘Ww
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FIGURE 175 (continued)
11b — conglomerates of Taman Peninsula, No.1249/232, PIN (Phanagorian(!? ) elephant);*
12а, 125 — Obitochnoe, Ukraine, No.24239, ZIN; 13a — Stavropol, No. 25284, ZIN
423
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(429)
428
430
The above-mentioned morphological changes in the teeth of the supposed
evolutionary sequence can be traced on the photographs of their longitudinal
sections (Figures 173, 174) and by casts and diagrams (Figure 175).
Table 91, which gives the morphometric indices of the teeth from some
"key'' burials in the Caucasus and the Russian Plain, is of practical
significance in stratigraphic correlations. *
The changes in tooth structure may be summarized as follows. From
the time elephants appeared in the region of the Black and Caspian seas to
the time of their extinction in the Lower Holocene, the maximum number
of enamel loops of Me increased from 16/15 to 25/19; the number of loops
over 10 cm of the worn or lateral surface increased by 4.1/4.2. Thickness
of the enamel wall decreased by 2.05/2.30 mm.
The apparently saltatory changes in the tooth structure of Quaternary
elephants, described so briefly in the preceding section, may be better
explained from the geological (taphonomic) rather than the biological
standpoint. The apparently isolated stages of morphogenesis of elephants
(and other animals) may correspond to the changes in the structure of
river networks and distribution, and in cycles of sedimentation which
produced the conditions necessary for the burial of bones. Evolutionary
inertia, which almost always operates in the evolutionary process, results
in the lag of adaptive changes behind rapidly changing environmental factors.
The accumulation of sediments depends, of course, on environmental
factors; however, we tend to regard the evolutionary changes described
as a continuous (not necessarily saltatory) accumulation of minor features,
terminated by evolutionary extinction.
Unfortunately, there are no data so far which would indicate that the
development of mammoth features in the elephant stock occurred in the
Caucasus at the same time as on the Russian Plain. Future comparisons
of the results of geological and paleontological studies will undoubtedly
reveal the truth.
Order PERISSODACTYLA
Very little work has been done on the phylogeny of perissodactyls, and
as yet there is no agreement among paleontologists about whether the
rhinoceroses of the Upper Pliocene and Anthropogene (Rhinoceros
etruscus, В. mercki, В. tichorhinus) represent a Single
phylorzenetic series. The pronounced morphological differences between
these stratigraphically separated species are less sharp in the transitional
stratigraphic stages. The skulls of the still poorly known
R. binagadensis from the Middle Pleistocene of the Apsheron Peninsula
(Binagady) differs from the Upper Pleistocene woolly rhinoceros in its
deeply concave intertemporal region, the smaller second horn and greater
inflation and callosity of the base of the first horn, the shorter postorbital
part of the skull, thin mastoid processes and small (spherical) occipital
condyles. The teeth of this species were considerably bigger (broader and
longer) than those of the woolly rhinoceros, and the aboral edge of the last
* In the construction of the table it was taken into account that the direct affinity of the Upper Pliocene
forms to the Quaternary forms has not been proved in all cases.
429
431
molar protruded far beyond the anterior edge of the orbit (Figure 176, 2).
The preorbital foramina are very small. The relatively narrow occipital
region of the skull of the Binagady species indicates that its head was held
in a more horizontal position than that of the woolly species. The Binagady
rhinoceros probably thrived more on tree branches than on grasses, as did
the woolly rhinoceros. The more slender metapodials, forearm bones and
tibia of the Binagady rhinoceros indicate that it was more lightly built and
was more active than the woolly rhinoceros. The abundance of this species
in the clearly xerophytic landscape of the Apsheron, very different from the
habitats of the woolly rhinoceros, attests to the distinct specific and even
generic identity of the Binagady form (see Dzhafarov, 1955). All the data
presented above do not preclude the possibility that the woolly rhinoceros
migrated from the north to eastern Transcaucasia and farther south into
southwest Asiain Upper Pleistocene times. This seems very likely, as it
is known that other large mammals like the primitive bull migrated in the
Upper Pleistocene fromthe north to the Apsheron Peninsula. Since the
remains of the woolly and early rhinoceroses in the Caucasus are
represented by fragmentary material, at present we can only give a map
of their occurrence (see Map 77), without going into a detailed analysis
of their origin and stratigraphic variation of morphological and phylogenetic
features.
FIGURE 176. Rhinoceros skulls; 1 — Rhinoceros tichorhinus, Upper Pleistocene,
Russian Plain, Vladimir, No.10699, ZIN; 2—Rh. binagadensis, Middle Pleistocene,
eastern Transcaucasia, Binagady, No. 24402, ZIN
430
The evolutionary trends of Elasmotherium (Е. caucasicum —
E. fischeri), known only from its Upper Pliocene to Middle Pleistocene
representatives, included a considerable decrease in the size of teeth and
development of plicated enamel plates (Borisyak, 1914).
The problem of the direct and indirect continuity of evolution of
morphological characters in horses from the Anthropogene of the Caucasus
and Russian Plain is as difficult as the problem of the evolution of
rhinoceroses. The great ability to migrate and the almost universal
adaptability to various types of habitat make it very difficult to solve the
432 problem of the origin of the east European and Caucasian horses in the
Quaternary satisfactorily. For example, it is still not clear whether
Przewalski's horse is an evolutionary successor of the local Upper
Pleistocene horse (broad-footed horse from the Upper Paleolithic of the
Don area), or whether it migrated from the south in relatively recent times
(i.e., Holocene). However, it is clear that local races of horses evolved
in the Caucasian Isthmus and the Russian Plain, replacing one another at
different geological periods. The general trend in horse evolution was
reflected in the development of hypsodonty and monodactylism, but the
details of the process varied from region to region.
Considering the data presented above, it is only possible at this stage
to speak of general stratigraphic and geographic changes in the post-Tertiary
horses of Russia, without relation to the evolution of any particular
species. As far as the Caucasian and east European material shows, the
evolution of horses proceeded as follows. The Upper Pliocene horses of
the Equus stenonis type from the Azov area were large, relatively
heavy, with large teeth, characterized by a relatively thin and much folded
enamel plate. The metapodia were neither highly specialized for fast
running in semideserts and steppes nor broad-footed and thus adapted
to the meadows and peat bogs of the taiga and tundra.
The horses of the Upper Pliocene-Lower Pleistocene of the
Е. sussenbornensis type (from the conglomerates of the Taman
Peninsula) had completely equuid teeth; their metapodia were lightly built,
indicating that this form was adapted for fast running on savannah and
steppes, which were regularly burnt during the summer. These horses
were bigger than the Quaternary horses (Vereshchagin, 1957a), but a
smaller horse occurred in the Middle Pleistocene of eastern Transcaucasia,
characterized by more antero-posteriorly flattened metapodial diaphyses.
Protuberances of the lower epiphysis ridge of the canon bone and
metacarpus are indexes that the osteochondrous type of pastern fixation
is somewhat less marked in the Binagady horse than in the Upper Pliocene
forms. The laterial ligaments, however, are stronger. The hoof phalanges
of this horse are relatively small, with a steeply rising angle anterior to
the wall, an adaptation to the xerophytic habitats and the rough ground of
this region. The fossil representatives of Upper Pleistocene and Holocene
horses of the Caucasus are still poorly known; however, horses of the
southern type, of "агу" contsitution (see Chapter III), undoubtedly inhabited
Transcaucasia. The Upper Pleistocene horse of the Russian Plain was
characterized by very massive legs and a brood foot, which can be seen
particularly well in the representatives from the Upper Paleolithic beds
at Kostenki on the Don (Figure 177).
431
(433)
`
За
ere *® Fea ea
-
FIGURE 177. Stratigraphic and geographic variation in the shape and supporting surface area
of the hoof phalanx of Eurasian horses. a — fore legs; b — hind legs. $ natural size
1а, 16 —-Equus caballus gmelini, Recent "tarpan", Russian Plain, No.521, ZIN;
2а, 25 — E. przewalskii, Recent Przewalski's horse, Dzungaria, No.17591, ZIN;
За, 3b — E, caballus subsp. (foss.), LowerHolocene, Novosibirskie Islands, №.4419, ZIN;
4a,4b —Е. caballus latipes, Upper Pleistocene (Upper Paleolithic) Voronezh, Kostenki XV,
No, 25183, ZIN; 5а, 55 —Е. caballus subsp., Middle Pleistocene, eastern Transcaucasia,
Binagady
432
434
These horses probably grazed on soft meadows, tundra and taiga swamps
on the floodplains of rivers. A similar structure of the hoof phalanx, though
not as wide, is characteristic of the horses of the extreme northeast of
Siberia, which probably lived on somewhat harder terrain under conditions
of thinner snow cover. *
TABLE 92. Stratigraphic and geographic variation of the supporting surface (mm 2) of hoof phalanges
of horses from the Anthropogene of Asia
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Note, Means in the numerator, observed ranges in the denominator.
The size of the area of the fore and hind leg hoof phalanges of different
horses and their interrelationships are given in Table 92. The data in
Table 92 show that in the southern and steppe varieties of horses the
anterior hoofs were relatively larger.
During the period of maximum cooling and development of mesophytic
landscapes, the broad-footed horse migrated far southward, probably to
the foothills of the Caucasus. There are reasons to believe that the
xerothermal stage and the extensive development of steppes on the Russian
Plain resulted in further divergence of these Upper Pleistocene horses in
the development of morphological characters related to forest and steppe
tarpans; confirmation is difficult, however, due to the absence of good
skeletal material of these so recently extinct forms (see Gromova, 1949;
Geptner, 1955).
Order ARTIODACTYLA
The large amount of work done on this order allows easy tracing of the
main evolutionary trends of some of the Caucasian forms.
* А.А. Sludskii (pers. comm, ) related the occurrence of wide hoofs on the fore legs of the northern and
Upper Pleistocene hoofed mammals to the need to dig through the snow in search of food.
433
435
Pigs of the group Sus scrofa — 5. vittatus appear in Eastern
Europe and the Caucasus for the first time in the Upper Pliocene, and
their main morphological features by then have already been eatablished.
Amon (1938) noted the extreme variation in skull morphology of the
Pliocene-Pleistocene pigs, which already inhabited vast areas of Eurasia
by the Lower Pleistocene. Therefore there are reasons for believing that
any stratigraphically younger pigs found in Russia are members of a
continuous evolutionary lineage.
The Taman boar (S. tamanensis) from the Upper Pliocene beds of
the Taman Peninsula differs from the younger Caucasian boars in its
larger size and, in mature males, a much wider lower jaw in the region of
the diastema (Vereshchagin, 1957a). The boar from the Middle Pleistocene
beds of the Apsheron Peninsula (S. apscheronicus) does not differ in
size from the Recent speciés, but the Apsheron species is characterized
by auditory bullae elongated downward and by the lachrymal bones of the
type intermediate between S. scrofa and S. vittatus (Burchak-
Abramovich and Dzhafarov, 1948). Widths of the lower jaw of Quaternary
boars measured behind the tusks are given in Table 93.
TABLE 93. Changes in size of boars by the dimensions of the lower jaw (in mm)
Width of lower
jaw behind tusks
Number of
specimens studied
Locality and geologic age Species
SUSE sero га ата | eas
Caucasus, Recent
Middle Don, 10-13thcenturiesA.D. | $. scrofa attila..... 3
Volga area, Tunguz Peninsula,
Middle Pleistocene
5 ЕЁ senofas: hiya. eee $ 1
Transcaucasia, Apsheron
Peninsula, Middle Pleistocene S.apscheronicus .
Ciscaucasia, Taman Peninsula,
Upper Pliocene
Я. сала ее
Note, Means in the numerator, observed ranges in the denominator.
A number of lower jaws and skulls of boars of the Anthropogene are
shown in Figure 178. The relative size of the bones of the axial and visceral
cranium has changed considerably since the Middle Pleistocene; however,
there have only been minor changes in tooth size since that time. The sizes
of M3 are identical in the Binagady and Recent Caucasian boar. However,
the heel portion of the tooth of the Recent species has become more complex
due to development of 1-2 columnar papillae on the heel (see Figure 178, 1, 4).
The abrupt saltatory decrease in the size of teeth, particularly noticeable
in M3, occurred only in the Neolithic in domesticated populations.
434
436
The remains which have been reliably identified as the red deer group
(Cervus elaphus s. lato) occur in the U.S.S.R. for the first time in
the beds of the Lowermost Anthropogene.* These deer evidently evolved
from some Pliocene Mediterranean forms. Pavlova (1926) has described
a skull and very simple unbranched antlers of a deer (Cervus
tschelekensis) from the Akchagyl (Upper Pliocene) beds of the
Transcaspian region. Isolated fragments of antlers (with the first outgrowth
somewhat above the crown) of a small variety of Pliocene deer occur in
the Lower Pleistocene conglomerates of the Taman Peninsula. Their
identification with the red deer group is not certain (Vereshchagin, 195%a);
though it seems possible that they are ancestral forms. The Lower
Anthropogene deer, known from the Tiraspol gravels (on the Dniester),
are characterized by antlers with two closely spaced low supraorbital
outgrowths and by the primitive, crownless type of the terminal branching.
FIGURE 178. Skulls and lower jaws of boars
la, lb — биз scrofa attila, Recent, Greater Caucasus, No.10485, ZIN; 2—S.scrofa attila,
10-13th centuries A.D., Lower Don, Sarkel; За, ЗЬ — 5. apscheronicus, Middle Pleistocene,
eastern Transcaucasia, No.26031, ZIN; 4—S.tamanensis, Upper Pliocene, Caucasus, Taman
Peninsula, No. 26000, ZIN
This type of "'tightly'' branching antler was remarkably stable, persisting
through the Anthropogene in Eastern Europe and the Caucasus. However,
the Middle and Upper Pleistocene deer of the Volga valley and areas farther
east are characterized by a ''looser" position of the first outgrowths on the
* Indications of earlier occurrences, as for example, in the Middle Pliocene of Transcaucasia (Bogachev,
1938c), are not certain. They probably refer to the deer of the genus Eucladocerus.
435
(437)
437
antler stem similar to that of the Recent Siberian and American wapiti.
Development of the fairly large antlers in the European noble deer began
probably in the Middle Pleistocene. A weakly-developed type of vestigial
crown is noted in some antlers washed out of the Khazar alluvium of the
Volga (Figure 179) and Don valleys, and the crown is well developed in
some antlers from the bitumens of the Apsheron Peninsula.
0 5 10 1§ 2025cm
a a oe)
FIGURE 179. Antlers of red deer (Cervus elaphus subsp.) from the Middle
Pleistocene alluvium of the proto-Volga valley. Kuibyshevy museum
The extensive descriptions of deer from the Binagady materials
(Alekperova, 1952) indicate that the Middle Pleistocene deer of eastern
Transcaucasia are not appreciably different in size from the Recent deer
of the Greater Caucasus. The Middle Pleistocene species is characterized
by lighter antlers. According to Akeperova's drawings, antlers with poorly-
developed and widely-spaced supraorbital outgrowths also occur in the
bitumen. This material is not sufficient for the study of the role of the
migrations and local speciation of the noble deer of Eastern Europe and the
Caucasus in the Anthropogene under the conditions of changing landscape
and ecology. Nevertheless it is clear that during the Quaternary in Eastern
Europe and the Caucasus there was а ''struggle'’ between the elaphus -
type (Cervus elaphus — with developed crown) and the wapiti-type
(С. canadensis — without crown and withwidely-spaced first outgrowths).
For the Caucasian population the development of this process can be traced
by using Dinnik's (1914a) descriptions and very complete coilections
as those of the Digorized cave in North Ossetia and the museum of the
Borzhomi nature reserve. The elaphus-type of antler is usually
suppressed in the Recent Caucasian deer, but in the Digorized collection,
which in 1948 consisted of 62 pairs and 121 single horns, 27% were
436
438
440
e laphus-type antlers (Figure 134) and 73 % wapiti-type.* The reliability of
these estimates is somewhat questionable due to the fact that many antlers in
this collection have an intermediate type of terminal branching, i.e.,
vestiges of a small crown.
Smirnov (1923-1924) pointed out that the Caucasian deer has decreased
in size since the Upper Paleolithic, his conclusions being based on the
occurrence of teeth in the Gvardzhilas cave. Indeed the teeth found there
by us and D.M. Tushabramishvili are of striking size, almost like those
of elks.
Since the collections contain mostly Quaternary deer material, it is
only possible to speak in greater detail of stratigraphic variation in antler
size. The antlers, and probably the animals themselves, gradually
increased in size from the Lower to the Upper Pleistocene, the largest
example occurring in the Upper Paleolithic of the Russian Plain (Kostenki,
near Voronezh). Later, in the Holocene, they somewhat decreased in size
again, though not as sharply as bison, for example; in the Middle Ages the
deer of the middle Don valley (Fortress Sarkel, near Tsimlyanskaya) were
still very large (Figure 180, 181). It is of interest that size change
in deer somewhat lags behind such changes in bison (see below).
The stock of particularly large individuals which colonized the Russian
Plain at the end of the Upper Pleistocene was probably derived from the
Caucasian population of deer which survived the Quaternary cool phases
under most favorable conditions. The migration of the Caucasian deer to
the north probably followed the Don and Volga valleys. In any event,
Flerov's (1952) statement that they attained their maximum size in the
Middle Pleistocene (Mindel-Riss) and then gradually began to decrease in
size is undoubtedly wrong.
The Recent Caucasian deer are characterized by body dimensions and
size of antlers intermediate between the West European and Asian-
American forms. This has been correctly noted in the first description
by Ogilby.
Geographic variation of the Recent deer C. elaphus maral inthe
Caucasus has not been studied in detail; variation is to a large extent
masked by individual and age variability.
According to Dinnik (1914a), the deer of the northwestern Caucasus
(Kuban area) are the largest, their size approaching that of Manchurian
wapiti. The size of the Caucasian deer decreases to the east and south,
and those inhabiting the foothills of Dagestan and the tugaic lower reaches
of the Terek and Sulak are particularly small. Both the fossil material
and the Recent populations are characterized by light horns, with only a
few (5-6) outgrowths, and almost always devoid of crown. The deer which
inhabited the Armenian Highlands and Talysh were also of relatively small
size. АП this is accounted for by the optimum conditions of the northwest —
abundance of food, vast areas of forests and mountains, etc. — aS compared
with the hot and dry east and south with their ancient civilizations.
Thus it can be seen that the picture of the development of the
morphological type and range of distribution of the red deer in Eastern
Europe and the Caucasus in the Cenozoic is extremely complex. This also
shows that some of the excellent paleozoogeographic analyses of Geptner and
Tsalkin (1947) can be confirmed or disproved only by means of additional
collecting and detailed study of paleontological materials.
* Dinnik's statement on the 500 pairs of horns at Digorized is somewhat exaggerated.
437
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FIGURE 181, Change in antler size in red deer of the Cervus elaphus group. One-half of the sum of
the diameters of the antler bases under crown. Black dots indicate means. Ordinate — frequency
Pliocene
Upper
~™ № & A
1— 15-19th centuries A.D., North Ossetia, Digorized cave; 2— 10-13th centuries A.D., Don valley,
Sarkel; 3 — 1st millennium B.C., Caucasus, burial, Makhachkala, Samtavro and Lake Sevan shores;
4 — Upper Pleistocene (Upper Paleolithic), Don valley, Kostenki XIV; 5 — Middle Pleistocene, bitumens of
Apsheron Peninsula, Binagady; 6 — Middle Pleistocene, alluvium of proto- Volga; 7 — Lower Pleistocene,
alluvium of proto-Dniester, Tiraspol; 8 — Upper Pliocene, Caucasus, Taman Peninsula
439
442
The phylogenetic lineage of the Recent elks of the genus Alces is
so far known only from the Middle Pleistocene. Nevertheless, there is no
doubt that Alces alces evolved at least in the Upper Pliocene, since
by the Middle Pleistocene the animals were already indistinguishable from
the Recent forms. The elk-like cervids of the side lineages which became
extinct inthe Lower Pleistocene, Tamanalces and Alces latifrons,
had, by the Upper Pliocene, developed some features of skull and antlers
similar to those of elks. These features developed, evidently, long before
the Anthropogene.
The adaptations of the Recent northern elk to life in the taiga swamps
and deep snow (long legs, large supporting surface of hoofs, brachydonty
and feeding on twigs) could have originated long before the elks invaded
the taiga and moss swamps.* Until recently, elks inhabited (in 1951) the
floodplain marshes of the middle Volga area. Thus, Flerov's (1950, 1952)
statements on the recent origin of these adaptations and on the feeding of
elks mainly on leaves and bark do not correspond with the facts.
It is known that for 7-8 months of the year elks feed mainly on hard
(frozen) twigs.
The skulls and antlers of Middle Pleistocene elks from the Khazar beds
of the Middle Volga region were already practically indistinguishable from
the Recent forms. The Caucasian elk, A. alces caucasicus, known
in the Caucasus since the Upper Pleistocene, was a relatively small animal
in the 18th century with clearly pronounced features of the European form.
The skull is relatively broad, particularly in the preorbital and frontal
regions; the nasal section is narrower than inthe northern elks. The
molars differ in their branchydonty with complicated wedge-shaped folds
on the labial surfaces of Pm® and Pm‘. In general, the morphological
features of the Caucasian elk are sufficiently well pronounced, which enables
one to date its appearance and beginning of isolation in the Caucasus as
at least the Middle Pleistocene.
The saiga (Saiga tatarica) is an example of considerable
morphological stability since the Middle Pleistocene. However, the Russian
and West European collections consist mostly of teeth and frontal-
occipital fragments with horncores. The oldest and probably most primitive
type of the frontal-horn fragment was collected in the Khazar beds
of the central Volga area. The skull of the Middle Pleistocene East
European saiga is relatively small, with absolutely straight, fairly
short and only slightly diverging horns (Figure 182, 4a, 4b), quite
similar to the skull of the saiga from the Binagady asphalt in
eastern Transcaucasia. While no different from the Khazar skull type in
the divergence of the horns, the horn stems of the Binagady skulls are
Upper Pleistocene skulls from the Khvalynsk beds of the Ural valley
(our collections, 1950) are characterized by greater divergence of the
horns, curvature of the horn stems in the frontal plane, massive structure
in some specimens and even greater length in other specimens
(Figure 182, 2a, 2b).
* It should be mentioned that all these features developed also in the absence of snow in Tertiary giraffes and
in some antelopes of the Mediterranean and Africa.
** Unfortunately, Alekperova's (1955) detailed description of the Binagady saiga does not help in drawing
conclusions,
440
(441)
Holocene
Pleistocene
Middle
FIGURE 182. Stratigraphic and geographic variation in structure of horn cores and
frontal-occipital fragments of saiga skulls. 1/5 natural size
Frontal and side view: la, 1b — Sai ga borealis Tscher., Lower Holocene, Yakutia,
Vilyui valley, No.13636, ZIN; 2a,2b—S.tatarica, Upper Pleistocene (Khvalynsk
age), alluvium of proto-Ural, No.24201 (10), ZIN; 3a,3b—S.tatarica b inagadensis,
Middle Pleistocene, bitumens of the Apsheron Peninsula, Binagady, №.22385, ZIN;
4a,4b —Saiga sp., Middle Pleistocene (khazar age), alluvium of proto-Volga, No.1084, ZIN
44]
FIGURE 183. Outline of hom stems and frontal-occipital section
of skull of the Recent saiga
1—Saiga tatarica, Volga-Ural.steppes, No. 7064, ZIN;
2—S,tatarica mongolica, northwestem Mongolia,
No.8465, ZIN
This type of frontal-occipital has persisted from the Upper Pleistocene
to Recent times in vast areas of Eastern Europe, central Asia and west
Siberia. In Yakutia, however, on the Vilyui and Lena rivers,
morphologically distinct populations of the northern saiga (S. borealis
Tscher.) are knownfrom the Upper Pleistocene and Lower Holocene,
characterized by the elongation of the region posterior to the horns, and
straight and long laterally compressed horns with weak divergence
(Figure 182, 1a, 1b).
The frontal-occipital region of the skulls in Recent saigas shows wide
variability; however, their skulls are practically indistinguishable from
the fragments of skulls from the central Urals (Figure 183, 1).
The geographic variation in Recent populations is probably related to
their occurrence on different types of pastures. In the upland semidesert
of Inner Mongolia, for example, an undernourished population has
developed, the subspecies S. tatarica mongolica Bann. (Figure 183, 2).
The sizes and frontal-occipital indexes of saigas are givenin Table 94.
443 Generally speaking, horn stems increased in length from the Middle
Pleistocene to the Holocene; the skull became wider between the orbits
and the region posterior to the horns shorter. At a later stage the horn
stems became shorter.
The size and proportions of the limb bones, the length of the tooth row
and the tooth structure of the Pleistocene saigas do not differ essentially
from the Recent forms. For example, the index of lateral compression
of the metapodial diaphyses, i.e., the ratio of anteroposterior diameter
442
444
of the diaphyses of metacarpus and metatarsus to the transversal diameter
(in the middle of the bone), varies in the Binagady saiga within the following
limits: metacarpus, 90-105% (5 specimens), metatarsus, 128-140%
(3 specimens); whereas the indexes of the Recent Volga-Ural saiga are 90-102
and 114-146 % respectively (5specimens). The jumping index, i.e., the length
ratio of tibia to femur, is 114.9% (3 specimens) in the Binagady saiga, and
114.2% (5 specimens) in the Recent species.
TABLE 94. Stratigraphic and geographic variation in size (in mm) and proportion (in %) of frontal-occipital
fragments of saigas*
Size index Length index
Horn stem " :
ии (circumference | Maximum (length of skull
Species, localities, geologic age at stem Базе | orbital skull section behind
measured ; } :
as % of its width horns as % of its
along chord i i
length) orbital width)
Northwestern Mongolia, Recent Saiga
: : : 99 62.7 106 62.0
tatarica mongolica, 1 specimen
Dzungaria, RecentSaiga tatarica, 127 76.7 125 47.5
ZISPECIIMENS) о 9 озноб oie ae) ново 124-130 76.7- 76.9 125-125 46.5-49.5
Central Don, 10-1316 centuries A.D. 139 718.0 62** 45
Saiga tatarica, 100specimens.. 110-165 71-91 130-144 44.2-417.0
Yakutia, Olenek, Lena Delta, Upper
Pleistocene, Saiga borealis, 146 70 122.0 56.1
Ре Boag 315 оо ooo devise Alb 139-153 66.7-73.4 117-127 55.0-57.2
Ural valley, Upper Pleistocene, Saiga 156.1 77 134.5 51.8
паважса , LOispecimensia №... 141-179 65.9-87.1 129-142 49.2-55.6
Apsheron Peninsula, Middle Pleistocene,
Saiga tatarica binagadensis, 135.5 75 59.9
2 SPECIMENS he... Rie Me Nemec ey ohne вл 135-136 74.3- 76.0 }
Volga valley, Middle Pleistocene, Saiga 115 82.6 57.5
SP LISPECIMEM es sterols) ile ial ein Nolte ils
Note. Means in the numerator, observed ranges in the denominator.
* Unpublished author's data, from ZIN collections. Selected measurernents giving significant differences.
** Based on 4 specimens.
In other words, the Binagady saigas were able to move as fast as the
Recent saigas; this feature also confirms to some extent the long-standing
stability of the morphological structure of this animal as mentioned above.
The stratigraphic variation of the Caucasian goats (Capra
caucasica, C. cylindricornis) is of particular interest in the study
of the origin and rates of evolution of the mountain mammals. However,
it is unfortunate that the bones found so far are mostly Upper Pleistocene
and Holocene.
443
The Upper Pleistocene (Upper Paleolithic) goats of the western Caucasus
differ from the Recent species in their larger size, which is shown by
measurements of the teeth and metapodia (Table 95). (The metapodia of
goats from the Gvardzhilas cave (Figure 184) are of remarkable size. )
FIGURE 184. Lower epiphyses of metacarpus (1-3) and metatarsus (5-7) of Caucasian
goats, Capra caucasica (foss.), from the Gvardzhilas cave, Upper Paleolithic;
metacarpal bone (4) of Recent 5-year-old C. caucasica
As far as the structure ofthe teeth and metapodiais concerned, noreliable
distinctive features are apparent inthe Upper Pleistocene and Lower Holocene
goats (Figure 97); the only exception is the somewhat greater eurosomality
(wide bones). In general, the main morphological and physiological
characters of the Caucasian goats — species adapted to mountainous terrain —
may be said to have developed by the Lower Anthropogene.
The geographic variation with longitude of the Recent Caucasian goats
is more clearly observable on the degree of specialization of the horns
of males, the structure of the horns becoming more complex from west
to east.
Four species of goats were identified in the Caucasus by the structure
of the horns, or, more correctly, horn sheaths: (from west to east)
1704 444
С. difan Mer бете лот Слезы са: са ара С. супа согииз
445 (see Dinnik, 1914a). Actually there are only two main forms: west and east
Caucasian.
TABLE 95. Stratigraphic variation in size of teeth and metapodia (in mm) of goats*
Upper Pleistocene, Sakazhia
Е Recent, west Caucasus
and Gvardzhilas caves :
Dimensions of teeth and metapodia
number number
mean range of of
specimens specimens
Anteroposterior diameter М?.......
Anteroposterior diameter M, ......
Transverse diameter of lower epiphysis
OigmMetacanpus ts.) «eNom
Transverse diameter of lower epiphysis
Gigmetatarsis; (1... -ieusriarela
* Measurements based on the largest individuals (8-12 years old for teeth and 5-7 years old for metapodia )
available in the ZIN collections.
The west Caucasian goat is more variable than the east Caucasian form.
The former is transitional from the central Caucasian C. caucasica
caucasica tothe west Caucasian C. caucasica severtzovi and
С. caucasica dinniki. However, transitional forms, probably hybrids
of the east Caucasian and Caucasian goats, occur in Svanetia, on the Upper
Ingur (Radde, 1899; Vereshchagin, 1938a).
The westernmost goats are characterized by massive and short horns,
sabre-like in profile, with slight divergence and ends slightly bent forward
and outward. In some specimens the horn sheaths display slight positive
coiling, 'homonimity'' of Nasonov (1923). The adjacent western goats
(C. caucasica severtzovi) are characterized by horns of predominatly
neutral ''ibex'' type with marked protuberances on the outer curved surface.
The ends of the horn sheaths diverge sharply; homonimity is slightly
developed, i.e., the right sheath coils to the right, and the left one to the
left. The central Caucasian goats (Guldenstaedt's turs) are characterized
by strong divergence of their horns, with ends slightly curved inward and
slight 'heteronimity'' of the sheaths, i.e., the right horncoils tothe left and
the left one to the right. The east Caucasian forms (Pallas or Dagestan tur)
have widely divergent horns; in old individuals the sheaths are bent
backwards and inwards, forming a low spiral. Slight heteronimity is also
observable. These forms of the geographic variation are illustrated in
Figure 185. In addition, the frontoparietal suture in west Caucasian goats
is straight, while its shape becomes more like an obtuse angle
in the skull of the east Caucasian populations.
If the original primitive form of the goat horns is the sabre-shaped
profile (ibex and wild goat), then it can be assumed that the horns of the
east Caucasian goat are phylogenetically the ''youngest. "
445
446
448
In view of the observed horizontal migrations and hybridization, the
origin of the ''latitudinal'' variation and morphological differentiation of
both forms of goat in the Caucasus can be accounted for by very high rates
of evolution of certain morphological characters. Taking into account the
Recent ecology of both species (Chapter III), there is no evidence which
would indicate that their isolation is the result of the glaciation of the
region. It is of interest to note that the genus Ovis, distributed over the
vast areas of Eurasia, represents another example of progressive
development of more complex horns in its range of distribution from west
to east.
FIGURE 185. Skulls and horns of goats of the Caucasus
1— Capra caucasica severtzovi, western Caucasus, No.12325, ZIN; 2—C. caucasica
caucasica, central Caucasus, No.697, ZIN; 3—C.cylindricornis, eastern Caucasus, No.6964, ZIN
The origin of the Caucasian goats and their closest generic affinities
are not quite clear. The ecology and morphology of the skull and horns of
Recent forms is similar to those of other goats of the ibex type inhabiting
Eurasia, being closest to the Alpine ibex (Capra ibex L.), yet it differs
strongly from the geographically closest Sinai ibex (C. sinaitica
Ehrenb. ) and Siberian ibex (С. sibirica Meyer). Even greater are the
morphologic differences in the structure of the skull and horns between
the Caucasian and the Spanish ibex (C. pyrenaica Schinz. ).
446
The two Caucasian forms — western and eastern — have, ofcourse, many
more morphological and ecological features in common with each other
than with any other named species of goat. For example, the lachrymal
bones and the portion of the skull posterior to the horns are identical in
the east and west Caucasian goats. Therefore the identification of
Cy bers se vert zo Меня. yaGCemtbicsc™auibaisana в. 'Cuvier and
CC. ibe x, si piciica Pall] asvonerspecies —C. ibex Li, аб was done
recently by Ellermannand Morrison-Scott (1951), may be regarded as an
unfounded extrapolation. * In spite of the fact that the Sinai and the Siberian
ibex resemble one another in the shape of the horns, the structure of the
lachrymal bone and the region of the skull posterior to the horns, it would
be erroneous to identify these species with the Alpine and west Caucasian
ibex. Moreover, for some reason С. caucasica is designated as a
species, whereas C.cylindricornis is designated as one of its
subspecies.
The morphologic distinction of the Caucasian goats from other Eurasian
forms attests to the isolation of their ancestors in the Caucasus in the
Middle Quaternary.
Among other cavicornia, the best-developed continuous series of
morphologic changes traceable since the Upper Pliocene is that of bison
(genus Bison).
The evolution of bison, like the evolution of elephants of the mammoth
lineage, took place in the vast areas of the Holarctic region as in Asia
and North America. However, the outcome and the features leading to
extinction of the Holocene populations of bisons on these continents were
entirely different (Vereshchagin, 1956).
Some American investigators, for example Skinner and Kaisen (1947),
regard the geologically distinct stages of the morphogenesis of bison on
the level of genus, which is probably incorrect.
In the Caucasus and the Russian Plain the phylogeny of bisons is known
from several geologic phases of accumulation of freshwater sediments
during the entire Quaternary era to the Recent: 1 — flat-horned small
Bison sp. (tamanensis N. Ver.) in the Apsheron age of western
Ciscaucasia (Taman Peninsula) and eastern Transcaucasia (Kabristan);
2 — considerably larger В. schoetensacki from the third terrace
of the Dniester (near Tiraspol); 3 — huge long-horned B. longicornis**
from the alluvium of the high terraces in Ciscaucasia and the Khazar
alluvium of the Volga and Ural river valleys; 4—B. priscus, asmaller
form from the drift and alluvial loams and gravels of the first and second
terraces in Ciscaucasia (post-Mousterian site, Il'skaya) and caves in
western Transcaucasia (Aurignacian site, Sakazhia, etc.), and also from
the Khvalynsk age and its equivalents on the Russian Plain (asphalt-
impregnated loams at Nizhnie Karamalki in Tataria and other localities);
5 — the sharply decreased size of the postglacial В. bonasus of the
historical epoch with two subspecies surviving to Recent times: the
Lithuanian B. bonasus bonasus andthe Caucasian B. bonasus
caucasicus (see Figures 186, 187).
Tsalkin (1955) has recently arrived at a similar conclusion.
** Рог the sake of nomenclatural consistency, the long-horned bison from the Khazar alluvium, which is
older than B. priscus Boj., must be regarded not as a subspecies of the latter but assigned the rank
of species.
447
(447)
--7
4
Holocene
Pleistocene
Pliocene
FIGURE 186, Stratigraphic and geographic variation in skulls and horn stems of bison of the Russian Plain
and the Caucasus. Contours of frontal surface of skulls and stems. 1/15 natural size
1— Bison bonasus caucasicus, 15-18th centuries A.D., North Ossetia, Digorized cave;
2—В. bonasus bonasus L., 10-13th centuries A.D., Don valley, Sarkel; 3— B. bonasus
bonasus L., 12th century A.D., Belorussia, Grodno; 4— В. priscus (aff. deminutus), Upper
Pleistocene (Upper Paleolithic), Transcaucasia, Sakazhia cave, un-numbered, Odessa University;
5—B.priscus deminutus, Upper Pleistocene, TatarA.S.S.R., Nizhnie Karamalki, No.1, Kazan
University; 6— B. longicornis, Middle Pleistocene (Khazar age), alluvium of proto-Volga, Stalingrad
Hydroelectric Station, No, 26134 (1), ZIN; 7-—B. schoetensacki, Lower Pleistocene, alluvium of proto-
Dniester, Tiraspol, Moscow Geological Survey Institute; 8 —В. cf. schoetensacki, Lower Pleistocene
Caucasus, Taman Peninsula, No.26009, ZIN; 9—B. sp. (tamanensis N. Ver.), Upper Pliocene,
Caucasus, Taman Peninsula, No.26010, ZIN
448
449
О ООО ООД ООО ООО ОО ОО ООО ООВ ОО ООО О ТО ИО О В Ри
28 29 30 HW 32 33 34 35 36 37 38 39 40 4 42 43 44 4 46 47 48 49 50mm
FIGURE 187. Stratigraphic and geographic variation in tooth size of bisons from the Russian Plain and
the Caucasus. Anteroposterior diameter МЗ on the level of alveola (M3—a; M3— b)
la,lb— Bison bonasus bonasus L., Belorussia, Belovezh, Recent; 2a,2b—B. bonasus
caucasicus, Caucasus, Recent; 3a,3b — В. priscus, Upper Pleistocene, western Transcaucasia,
Gvardzhilas and Sakazhia caves; 4a,4b — В. priscus, Upper Pleistocene, Trans-Kuban Plain, П'зкауа
site; 5a,5b — В. longicornis, Middle Pleistocene (Khazar age), alluvium of proto-Volga, Stalingrad
Hydroelectric Station; ба, 6b —B.cf. schoetensacki, Lower Pliocene, alluvium of proto-Dniester,
Tiraspol; 7—B. sp. (tamanensis N. Ver.), Upper Pliocene, Caucasus, Taman Peninsula
Even the Upper Pliocene fragments of skulls and long bones give an
idea of the stabilized morphological features of the genus which evolved in
the Pleistocene. The rest of the lineage, as far as the available material
shows, was characterized by evolution of quantitative (size, length, volume)
rather than qualitative characters. The isolated teeth and long bones of
the long-horned bison from the Khazar alluvium of the Volga are essentially
449
only larger ''copies'' of the Lower Pleistocene bisons from the Tiraspol
450 gravels and the Upper Pliocene bisons from the Taman conglomerates.
Nevertheless even minor changes in the skeleton often attest to pronounced
changes in the external environment.
TABLE 96. Dimensions (in mm) of skull fragments of Pleistocene bison*
Length of horn Anteroposterior Postorbital
stem proper diameter of base of forehead width
Locality and geologic stage along the chord horn stem proper (number of
(number of зрес!- |(number of specimens specimens in
mens in parentheses)| —_in parentheses) parentheses )
Caucasus, Digorized cave, 16-18th 128(18) 60.1 (63) 223 (27)
GENT UIGIES, Al De ее Е 95-185 44-90 186-263
с с
И о cave, Upper 210 85 В
РАНЕ ов ана
.5.5.К., Ni i i, U
Tatar A.S.S , Nizhnie Karamalki, Upper 270 87 291
PleIstOCene! a ius. а seme зазье иен чене до
Volga area, Stalingrad Hydroelectric Station 350 (7) 120 (8) 328 (7)
localities, Middle Pleistocene ....... 371-560 105-140 301-352
308 (8) 106 (8)
davi i i ys ЗАУР Е =
Moldavia, Tiraspol, Lower Pleistocene 290-340 85-120
Caucasus, Taman Peninsula, Upper 210-260 80- 90 140
PIOCENE ss einen Sekt om aoe (5) (2)
Note. Means in the numerator, observed ranges in the denominator.
* The table is based on unpublished work of the author: measurements of male and female skulls in
Digorized cave, and of mature Pleistocene males in the collections of the Kazan Geological Museum,
Moscow Geologic Survey and a number of muzeums in cities on the Volga, the Odessa Paleontological
Museum, and ZIN.
For stratigraphic purposes, the dimensions of the skull fragments
(Table 96), in addition to Figures 186 and 187, are of importance,
particularly in the study of sexual dimorphism, age and individual
variability.
Studies of the morphogenesis of bisons in the Caucasus and adjacent
areas suggest that these hollow-horned forms evolved here during the
Anthropogene, and relict populations have survived in the mountain forests
of the regions into Recent times.
The history of the primitive bull (genus Bos) developed along different
lines, as in the Pleistocene the Caucasus was inhabited by forms of
diverse origins (Burchak-Abramovich, 1957).
In a discussion of the morphological evolution of Caucasian and east
European mammals in both time and space, it becomes clear that the
Quaternary index forms (carnivores, proboscideans and ungulates) evolved
450
451
452
in the Caucasian Isthmus in а manner analogous to the vast areas of northern
Eurasia. Some features of their evolution are identical with those described
by Hooijer (1949) for the southeast Asian mammals. As arule, most of the
animal species increased in size during the Pleistocene, followed by strong
decrease in size at the end of the Pleistocene as if Bergmann's rule
applied to distribution in time.
From study of the morphological changes in mammals during the
Anthropogene one concludes that most of the evolution of the Quaternary forms
took place some time before the Quaternary. Lower Pleistocene and even Upper
Pleistocene mammals (carnivores, rodents, proboscideans and ungulates)
show the characteristic features of Upper Pleistocene and Holocene forms
at the rank of genus, and even subgenus and species; thus we recognize
the fox from the group of the red fox, hamsters of subgenera Cricetus
and Mesocricetus, deer of the group Cervus elaphus, etc.
Since the onset of the Anthropogene, successive stages of related
mammals (in most cases) differ mostly in questions of proportion and in
relatively minor adaptations (Severtsov, 1939). For example the red deer of
the Upper Paleolithic and the historical epoch of the Don area is an enlarged
copy of the Lower Pleistocene deer of the Tiraspol gravel. There is an
apparent absence of qualitative saltatory changes in the Quaternary history of
mammals. This probably requires arevision of the criteria for the identification
of species. It is due to these facts that some Russian paleontologists, among
them Flerov (1952) and Gromov (1948), are mistaken intheir interpretations
about the geologically young age of certain mammals (reindeer, elk) and their
morphologic-physiologic adaptations to the conditions of taiga and tundra.
Their assumptions only indicate lack of understanding of the genesis
and evolution of the landscape of the northern part of the Holarctic region. *
To us it is beyond doubt that the Anthropogene was the period of development
of distribution areas and ecological assemblages of species which have
reached the final stages of their evolution.
The Anthropogene mammals mostly exploited ecological (physiological)
plasticity, developed since the Pliocene in response to changes in the
landscape behind which the morphological evolution often lagged. In other
cases animals became extinct or migrated into other areas.
The activity of man may possibly be partly responsible for the striking
interruption of the evolution of many Quaternary species and their sudden
extinction.
Among other general phenomena is the variability in the rates of
speciation and the heterogeneity of characters in different phylogenetic
lineages, sometimes even in taxonomically closely-related forms. We may
also point out the conservative nature of the skull characters of foxes,
as compared with the characters of the corsac fox, the slower rates of
increase and decrease in size of deer and elks as compared with bisons, etc.
* Most recent geological and paleontological studies in northem Siberia, particularly in Yakutia,
have shown that during the Cenozoic this country underwent considerable climatic changes together
with changes in landscape structure and fauna. The changes were similar to those which occurred in
the Mediterranean. Consequently, occurrences of Lower Quaternary reindeer, elks and saigas may be
anticipated.
451
In а number of cases Ц is not clear whether we are concerned with minor
adaptations to environment and initial stages of divergence, as has been
demonstrated by Vinogradov (1946) for the rodents, or whether we are
dealing with continuously developing characters within the framework of
a general evolutionary trend. For example, it is difficult to decide without
additional studies whether the increase in the length of the tibia and
increased inobility of the Recent jerboas and gerbils of the Apsheron
Peninsula, as compared with the Pleistocene forms, is the result of
development of xerophytic (desert) landscapes, or whether it is the
product of evolution of the organisms in a certain direction. The latter
explanation is presumably favored by examples of specialization in the teeth
of wolves, corsac foxes and elephants.
The main forces behind changes in morphology during the Anthropogene
were, of course, marked environmental and climatic changes.
Minor responses to ecologic variation include the independent
development of relict forest-mountain forms, which led to far-reaching
divergence and conservatism (Caucasian voles, goats), and also the rapidly~
developing initialstages of speciation which led to the establishment of
local ecological varieties (moles, foxes, common field mice, water voles,
etc. ).
The Caucasian populations of Eurasian species formed, as arule, local
subspecies from which not only their origin and the antiquity of their areas
of distribution can be determined, but also the rates of morphological
divergence.
The geographic changes in Recent mammals of the Caucasian Isthmus
are mainly expressed in decrease in size and brightening of the coloration
towards the southeast (foxes, snow voles), increase in body size with altitude
(susliks), and more complex structure of the horns of cavicorn species
towards the east of their area of distribution (Caucasian goats). These
examples are often only small-scale reflections of similar phenomena
observable over the vast expanses of Eurasia, and contribute to the
understanding of the origin and history of colonization of the ecological
niches in this area. It becomes increasingly clear that in the process of
development of this Anthropogene mammal fauna the main role was played
by the immigrations and the fluctuations in abundance and extinctions
of organisms, whereas the divergence and morphogenesis in general
were of secondary significance.
452
453 РатЁ Three
GEOGRAPHIC ZONATION AND LATEST
EVOLUTIONARY TRENDS OF MAMMALIAN
FAUNA OF THE CAUCASIAN ISTHMUS
455 Chapter V
ANALYSIS OF HOLOCENE MAMMALIAN FAUNA
AND SCHEME OF ZONATION
Through our studies of the origin and ecology of a number of Caucasian
mammals (Chapter III) it is possible to identify the principal ecological
and geographic groupings of those species whose ranges lie in more or
less clearly defined regions of diverse origins and ages. A study of the
distribution of the other Holocene animal species and an evaluation of their
origins and histories in the Caucasus would be a considerable contribution
to the zoogeography of this region. The zonation of the area into mammalian
geographic districts is necessaryin order to understand the regularities
in faunal evolution as they relate to changing landforms and to gain some
perspectives for the planned restoration of the fauna.
However, since the ranges of individual species and the composition of
ecological assemblages and faunal complexes are not constant, these
zoogeographic boundaries cannot be regarded as permanent. The more the
landscapes and the boundaries between landscape zones change, the greater
is the rate of evolution; and the less adaptable the animals are to these
changing environmental conditions, the greater is the rate of change within
the ecological and faunal complexes. Therefore, a zonation, particularly
on the scale of subregions or provinces, can be established only for
relatively short periods of geologic time. This fact is not always
recognized by zoogeographers. The concept of geographic boundaries of
a range of a ''faunal type,'' a ''faunal complex, '' etc. is only a convention.
Such a boundary only holds for an individual species on a specific date.
At any given moment in geologic time, it can only establish an approximate
coincidence of a number of species which are similar in their ecological
characteristics.
The schemes of zoogeographic zonation and zoogeographic categories
based upon characteristics of the species complexes presented by
contemporary zoogeographers are usually based on relict species, which
have survived during the historical epoch under various anthropogenic
influences. Only tentative, inadequate attempts have been made to employ
historical data on the occurrence of some extinct forms in the near past
and to infer from this history the dynamics of the ranges of individual
species.
The error in this approach lies in the exclusion of those species which
have recently become extinct, particularly those exterminated through
456 human activity. This omission rules out any possibility of gaining
perspective on the restoration of the fauna, which is the practical goal of
zoogeographic studies.
453
457
From the second half of the 19th century, almost every investigator who,
in one way or another, was concerned with the zoogeographic division of
Eurasia turned his attention to the origin of the fauna of the Caucasian
Isthmus and its zoogeographic zonation. Wallace (1876) divided the Isthmus
longitudinally along the Bolshoi Range, including Ciscaucasia in the European
region, and Transcaucasia in the Mediterranean region.
Severtsov's scheme (1877) was exceedingly simple and very nearly
correct. He assigned the northern Caucasus, western Transcaucasia and
the Caucasian Range to the Mediterranean region, the rest of Transcaucasia
to the western Asian region, and northeastern Ciscaucasia to the central
Asian region. This zonation reflected the heterogeneous origin of the fauna
of the Isthmus.
W. and Ph. Sclater (1899) combined the Caucasian Isthmus with Asia
Minor, assigning both areas to the Europaeo-Asian region.
It was Satunin who most clearly recognized the heterogeneous origin of
the Recent fauna of the Caucasian Isthmus in his papers of 1901d, 1903b,
1904-1906, 1909-1910, 1912b, and 1913.
His main hypotheses were the monophyletic origin of species and their
unvarying migrational tendency. * He evidently assumed that the focus of
speciation lay somewhere to the south of Transcaucasia and from there,
as from a cornucopia, the species spilled forth in northward migrations via
the Balkans and the Caucasus — the highly mobile species and those less
mobile. Satunin was the first Russian zoologist to attempt to synthesize the
history of faunal development on the Caucasus with the geological data
available in his time on the Tertiary and post-Tertiary seas, straits and
glaciations. He reasoned his migrational hypothesis from his belief that
evolution proceeds very slowly in mammals, and that the Caucasian Isthmus
is geologically much younger than the adjacent areas of southwest Asia and
southern Russia. Consequently, he concluded that no independent speciation
could take place on the Caucasus. According to Satunin, the Manych strait
and the Aral-Caspian basin in the northeast constituted barriers to mammal
migrations from the north to the Caucasus.
Satunin made an interesting attempt in 1901 to establish the geologic
age of mammals which migrated to the Caucasus. He considered that
Microtus arvalis and Arvicola amphibius already lived in the
Caucasus in glacialtime. After the disappearance of the Manych strait
"only Putorius foetidus, Vulpes corsac, Cricetus vulgaris,
Sminthus subtilis, Castor fiber, Allactaga satrews,
Lepus europaeus and Saiga tatarica migrated to the Caucasus
from the north.
"In addition to Cricetus vulgaris, mentioned above, Mus
agrarius and М. minutus _ should be included among the most recent
immigrants from the north."
Later (1900-1910) Satunin examined the origin of the fauna of the entire
Caucasian area and divided it into the northern Caucasus, the Range and
Transcaucasia. He defined nine subdistricts in these three major districts.
His work gave particular emphasis to the great antiquity and independent
* We are in agreement with Pidoplichko (1953) that local development of a number of mammalian species,
assemblages and complexes was a significant factor in the evolution of the Palaearctic, particularly the
eastern Mediterranean fauna. His point, however, by no means excludes the possibility suggested by
paleontological data of a dispersion of species and faunas during the Cenozoic, particularly in the
Quatemary.
454
origin of the alpine fauna and to the occurrence of immigrants in the forest
zone and on mountain slopes. He further hypothesized that the ancestors
of the mountain fauna of the Caucasus migrated from the mountains of
southwest and central Asia. He pointed out that the absence of ptarmigan
and blue hare in the highlands indicated that the arctic fauna never reached
the Caucasus, that the mountain fauna of southwest Asia is autochthonous
in nature, that there were no African elements in the Transcaucasian fauna,
and that the fauna remained purely Asian in character.
He accounted for the emergence of ''Aral-Caspian'' fauna in the Caucasus
by the glacial advance and the flooding of the Caspian steppes, and for the
isolation of the Aralik section in the Araks valley by vulcanism in the
area of the southeastern spurs of the Zangezur Range. He regarded the
fauna of the wooded parts of Talysh as the northwesternmost sector of the
Indian fauna.
Satunin briefly summarizes the general characteristics of the fauna of
the Isthmus in this way: ''In this region the following occurrences can be
observed — an ancient autochthonous fauna of the Greater Caucasus, a relict
Mediterranean fauna in western Transcaucasia, the original, endemic fauna
of the uplands of eastern Asia Minor, the northwesternmost advance of the
Indian fauna and, finally, the faunas which migrated from the steppes of
southern Russia and the deserts of central Asia."
Satunin's zoogeographical studies culminated in his work ''On
Zoogeographical Districts of the Caucasian Territory'' (1912b). He
subdivided the territory into five subregions and eleven districts. His
zoogeographic scheme of the Caucasus has been well accepted.
He saw his main achievement in the fact that his zonation almost
completely coincided with the phytogeographic divisions of the Caucasus.
The coincidence is not fortuitous — Satunin travelled widely over the
territory and placed great importance upon direct observation. He held that
the fauna does not merely depend upon a specific landscape, but forms an
integral part of it.
Satunin's compilations of mammals for various districts of the Caucasus
are extremely accurate and comprehensive. The interpretation of the data,
however, was hindered by prevailing notitions and by too little knowledge of
the ranges and faunas of adjacent territories.
Some of Satunin's errors were repeated in the last work cited: the
zoogeographic scheme remained ''geomorphological'' in nature, based on
the features of the ''phytolandscapes'' rather than on the actual distribution
areas of species and complexes. The boundaries either followed the
mountain divides or were drawn along the foothills, and in the south of the
country they were drawn from inference. The migration route of the Aral-
Caspian fauna was still considered to be through eastern Transcaucasia
along the ravine and valley of the Araks to the foothills of the Ararat and
Alagaz, although the Lesnoi district of eastern Transcaucasia was mapped
as a remarkable wedge-like extension, reaching to Artvin in the west and
including, on the northeast, half of the woodless interior of Dagestan.
Dinnik (1911), among other Caucasian faunologists, recognized the
predominantly southern aspects of the mammalian fauna of the Caucasus.
He pointed out the occurrences of three species of European, five species
of Eastern European and thirteen species of Asian mammals in the steppes
of the northern Caucasus. According to Dinnik, the mountain species
migrated to the Caucasus from the south, mainly from Asia Minor.
455
458
459
In a criticism of Satunin's work published 27 years later, Lyaister (1931)
made the point that 85% of the mammals in the Aralik section of the Araks
valley, which Satunin included in the steppe district of-eastern
Transcaucasia, are shared with adjacent Iran, and that only 41% are also
common to the Transcaspian region.
Actually only two species of the Aral-Caspian fauna are found in the
middle Araks valley: small five-toed jerboa and manul, and the ancient
links of these species withthe Transcaspian region were through northern
Iran, rather than through eastern Transcaucasia and Ciscaucasia.
Lyaister, like Satunin, in holding to the theory that most of the animals
characteristic of the Armenian Highlands migrated from Iran, was only
paying lip service to the unfounded hypothesis of ''migrationism.'' The
existence of dry land in this area since the Oligocene means that the Recent
Lesser Caucasus uplands can be regarded as a marginal part of the ancient
focus of speciation in southwest Asia.
Menzbir (1934), who was also well acquainted with paleontological
material, included all the plains of western Ciscaucasia in the Pontian-
Barabanian subprovince of the European-Siberian subregion, eastern
Ciscaucasia in the Kirgiz subprovince of the central Asian subregion
and the Caucasus and Transcaucasia in the Caucasian subprovince of the
eastern Mediterranean province of the Mediterranean subregion of the
Palaearctic region.
The northern boundary of the Caucasian subprovince defined by Menzbir
is in the foothills of the northern Caucasus on a line from Novorossiisk
оп the Black Sea coast to Khachmas on the Caspian. The southern boundary
is drawn from Batumito Yerevan around southernTalysh. The faunaof this
subprovince, which includes the southern Crimea, is mixed Europaeo-Asian.
According to Menzbir, the Caucasian fauna was affected by two faunal
centers —the Mediterranean and the central Asian.
The boundaries of Menzbir's subprovinces coincide almost exactly with
the broadly defined limits of the climatic and phytolandscape zones of
the Recent. Clearly, Menzbir's work simply gives the faunal characteristics
of the landscape zones of the Isthmus.
Puzanov's treatise (1938a) of the fauna and nature of the Caucasian
Isthmus is based on broad knowledge of geographic facts and presents a
concise, lucid discussion.
He places all of Transcaucasia in the Mediterranean subregion, grouping
it with the Greek archipelago, Asia Minor and mountainous Crimea in the
eastern Mediterranean province. The northern slopes of El'brus and Talysh
are included in the Asterabad province, adjacent to the central Asian
subregion.
Puzanov describes the Greater Caucasus as a ''complex faunal knot in
which Mediterranean forms predominate to the south, European forms to
the north, and central Asian forms to the east. The nucleus of the fauna,
however, is made up of an endemic alpine fauna of great antiquity."
Kuznetsov (1949, 1950), who was familiar with the extensive data
compiled in Soviet time, proposed a special geographic zonation of the
mammalian fauna of the Caucasian Isthmus. In his scheme of the Palaearctic
region, the plains of Ciscaucasia and the eastern Transcaucasian plains
are included in the zone of deserts and steppes, the foothills of Ciscaucasia
in the transitional forest-steppe zone, and the largest part of Transcaucasia
in the Mediterranean province of the subregion of northern forests.
456
His delineations of six Caucasian districts and nine subdistricts were
drawn on the basis of characteristic features of a fauna comprised of
113 species and on Bush's descriptions (1935) of the phytolandscape zones.
Both of Kuznetsov's papers were hastily published and contain much
erroneous data and many conflicting conclusions. Although he was correct
in his understanding and interpretation of the Mediterranean subregion,
which includes almost all of the Caucasian Isthmus, his treatment of the
pattern of faunal development is inadequate, as can be seen in the following
quotation: ''The third group of species of Caucasian mammalian fauna
consists of forms of definite Mediterranean (! ? — М. У. ) origin.'' From
this statement it would follow that the Caucasian, i.e., Mediterranean,
forms originate within themselves. His discussion of the Armenian Highland
is also vague, since the same area is referred to as the Armenian forest-
upland region and the upland Armenian region. Talysh is incorrectly
described as a forest-mountain area, although Satunin clearly distinguished
both mountain-forest Indian types and upland-steppe Iranian types in the
Talysh fauna.
Kuznetsov omitted the following forms from his list of species: the
vespertilio bats (У. ognevi and У. bobrinskii), pocketed bat, Asia
Minor field mouse, Caucasian snow vole, saiga, goitered gazelle, bison,
and recently extinct forms, i.e., elk, kulan, tarpan. There are also many
errors in his conclusions on the origin of the fauna, in the faunal
characterizations of the districts, particularly in the zoning of the Caucasian
district, and in the descriptions of boundaries.
The Azerbaidzhan district is an example. Its boundaries are described
by Kuznetsov (1950) as follows: ''From the town of Astara on the Iranian-
Soviet international boundary near the eastern end (? —N.V.) of the Talysh
Range, the boundary follows the foothills ridge (! ? —N.V.) in the southern
marginal area of the Mugan, Mil'skaya and Karabakh steppes to Lake Sevan
(! —N.V.) and farther through the town of Karaklis, south of Tbilisi, to the
Surami pass. Here the boundary turns sharply to the east through the towns
of Gori, Mtskheti, Lagodekhi, Zakataly and Nukha toward the eastern end
of the Main Range and over the Kuban lowlands toward the Caspian coast"
(p.157). This description creates complete confusion since, according
to it, the subdistrict of steppes and deserts includes both the Lenkoran,
Alazan-Agrichai and Khachmas lowlands with almost subtropical forests
and the northern slopes of the Shakh-Dag and Trialet ranges, surrounding
Lake Sevan, with beech—hornbeam forests.
The marginal sections of the Kura-Araks lowlands were never covered
by the waters of the Aral-Caspian basin in the Quaternary, rather than
being covered ''manytimes.'' And, therefore, rather than being a young
fauna, as Kuznetsov maintains, the fauna is in fact so ancient that it contains
such elements as Asia Minor hamster and mole (! ).
In the mammals listed for this district, which Kuznetsov described as
a plain with wormwood-steppes and floodplain forests, he included such
species as the common and lesser shrews, harvest mouse, forest vole
(1? — М. \.), pine vole and common vole, which definitely do not occur
in the Kura-Araks, Lenkoran and the greater part of the Alazan-Agrichai
lowlands. And such characteristic species as pine marten, serotine
vespertilio, Ognev's vespertilio, tiger polecat, panther and red deer are
not included.
457
460
However, Kuznetsov's interpretation of the Mediterranean subregion
is more accurate than interpretations of earlier authors.
In recent years the refinement of the ranges of rodents in western
Transcaucasia has engaged the attention of Shidlovskii (1940c, 1941b,
1945, 1947, 1948, 1950, 1951). He established the great diversity of the
faunal influences on the rodent fauna of Georgia and refined the district
boundaries laid down by Satunin (Shidlovskii, 1941b). In distinguishing
some species of rodents as immigrants to Georgia from the west, east
and south and others as ancient relicts, he presents a correct zoogeographic
position. Shidlovskii attempted to establish the geologic age and history
of the ranges of some of the species which comprise the rodent fauna of
Georgia. For example, he places the migration of the Asia Minor hamster
to the Caucasian Isthmus at the end of the Miocene- Lower Pliocene, and
relates the discontinuity in the range of Promethean vole on the Greater
Caucasus and in Adzharia to the development of xerothermal conditions on
the Dzirul'skii massif in postglacial time. Shidlovskii, at variance with
Satunin, proposed designating the alpine zone of the Greater Caucasus as
an independent zoogeographic unit, based on the presence of Promethean
vole and Caucasian birch mouse.
Bobrinskii (1951) relates the fauna of the main part of Ciscaucasia to
the fauna of the European-Kazakhstan steppes, the fauna of the northeastern
section of Ciscaucasia to the fauna of the Aral-Caspian deserts, and the
fauna of eastern Transcaucasia to the faunas of the Aral-Caspian deserts
and the European-Kazakhstan steppes. The entire forest-mountain fauna
of the Greater Caucasus, the northern slopes of the Lesser Caucasus and
the eastern slopes of Talysh he relates to the fauna of the European
broadleaf forests, and that of the alpine zone of the Bolshoi Range and
Lesser Caucasus to the mountain fauna, like those of the Kopet-Dag, Tien
Shan, Altai and the eastern Siberian ranges. Bobrinskii, quoting Satunin
and making use of his map, repeats his error on the predominance of the
Aral-Caspian fauna in Transcaucasia. His description of the fauna of the
Lesser Caucasus uplands also abounds in error (p.314), such as his
statements on the absence of deer, pine marten, mole and Bezoar goat
from that area.
Bobrinskii discussed the origin of the Caucasian fauna on the basis of
modern geological data. He maintained that the migration of European
animals to the Caucasus mainly followed the southwestern route via the
Balkans and Asia Minor, and that the extension of the ranges of the European
common vole and snow vole to the Caucasus was only possible through
large-scale migrations.
In addition to these disputable statements, he made the strange statement
that the isolation of Promethean vole, Caucasian black grouse, goat and
Snow partridge began at the time when the Main Range was still an island,
when in fact the Caucasus was not an island in the Oligocene and in the
Lower and Upper Miocene.
A very detailed zonation of the Armenian territory into ecological and
landscape districts was made by Dal' (1954a), who compiled lists of land
vertebrates from various natural, mainly phytolandscape, zones of the
Armenian Highland and the middle Araks valley. The fact that he employs
four vertebrate classes concurrently in his zoogeographic analysis of the
region, and that he adheres to the theory of a Eurasian ''faunal type",
458
461
creates considerable difficulty in ascertaining the origin and composition
of the mammalian fauna of Transcaucasia, even for the author. In his
zonation of Transcaucasia, Dal', inthe main, followed Kuznetsov's
(1949) scheme.
This brief review of zoogeographic concepts of the Caucasian mammalian
fauna of the Recent indicates, as did our paleontological data (Chapter ПТ),
that the ancient local focus of speciation and of faunal development in the
mountainous parts of the Isthmus was under strong influence from
mammalian complexes of adjacent areas.
In order to make a correct evaluation of these faunal influences and their
distribution in time and space, it is necessary to discuss the age and origin
of the local Caucasian and the neighboring faunal complexes. It is
particularly important to understand the origin and the nature of the
Mediterranean complexes — a subject on which there is no general
agreement. Paleogeographers consider the Mediterranean littoral as the
relict basin of the Tethys sea, or, at least, of the Sarmatian sea which
extended from the Atlantic to the Transcaspian region. In the climatological
sense, the Mediterranean is regarded as comprising southern Europe,
north Africa, part of southwest Asia and even part of central Asia (see
Berg, 1938).
Geobotanists Alekhin (1938) and Vul'f (1944) include the coastal areas
of the Mediterranean and Black seas in the Mediterranean region of the
Holarctic north Africa, Spain, Italy, Greece, Asia Minor and the Crimea.
And Maleev (1946) and Grossgeim (1936, 1948) have repeatedly pointed out
the genetic affinities, if not the identity, of Caucasian formations of maquis,
garigue, phrygana and broadleaf forest with similar formations in the
western Mediterranean, clearly establishing them in their view as one
unit. Lavrenko recently published the opinion (1958) that the forest region
of the Caucasus is similar in the phytogeographic sense to the broadleaf
forests of Europe in its origin and ecology (climate), but not to the
Mediterranean. Earlier zoogeographers often considered the Mediterranean
region to include the coastal areas of the Mediterranean Sea, Asia Minor,
northern Arabia and Transcaucasia (Wallace, the Sclaters, Severtsov,
Kobelt).
Modern ornithologists like Serebrovskii (1928) regard southwest Asia
(Sumer) as the area where the faunal elements of Africa, central Asia,
India and Europe converge. According to Shtegman (1938)the Mediterranean
type of fauna, apart from being found on the Mediterranean coasts, can
also be found in the Sumer of Serebrovskii, but the fauna of Transcaucasia
and the southern Crimea is characterized only by a tinge of the
Mediterranean type.
Kashkarov and Korovin (1931) observed the Mediterranean influences
in the flora and fauna of even the middle life-zones in Tien Shan. There
is a recent tendency to regard the Caucasian and Hyrcanian fauna as a part
of the province of broadleaf forests of Europe (Rustamov, 1945; Kuznetsov,
1950; Bobrinskii, 1951).
Geptner (1939, 1940, 1945) developed the idea that the centers of origin —
he named four — of the mammalian desert-steppe fauna of the Palearctic
all lay far to the south. He opposes (1945) the notions of an endemic origin
of Mediterranean fauna and of a Mediterranean region on the assumption
that there were no ancient and specific mammalian forms in the
459
462
Mediterranean, maintaining that the coasts of north Africa and southern
Europe are inhabited only by immigrants from the north and the south.
These assumptions are invalid because no true comparison can be made
between the landscapes and the diversity of biotopes in the deserts and in
the Mediterranean.
Animal immigration to the Mediterranean was always more open than
to the Sahara and Kara-Kum, but this does not detract from the importance
of the Mediterranean as an ancient independent center of speciation and
faunal evolution.
A correct interpretation of the concept ''Mediterranean region" (and a
resulting understanding of the Mediterranean type of fauna) can only be
reached through an analysis of the natural changes which occurred in the
Cenozoic in the Mediterranean geosyncline.
It is known that the Sarmatian sea, which extended from Gibraltar to
the Transcaspian area, repeatedly formed in later — Pliocene and
Pleistocene — time a chain of isolated basins, which were sometimes
interconnected during periods of transgressions. Their separate coastal
areas evolved under varying conditions. The Aral and northern Caspian
coasts because of their relief and greater continentality early became
deserts, which excludes them from the unique system of Mediterranean
zonation.
The other coastal areas of this chain of Cenozoic basins, including the
Caucasus, inherited a number of unique, primitive features of landscapes,
flora and fauna, which originated in the Miocene. Therefore, the southern
areas of Europe (including the southern Ukraine and the Crimea), north
Africa, Asia Minor, the Caucasian Isthmus (excluding the Manych area)
and the Talysh and El'brus ranges should be included by virtue of their
paleogeographic identity in the Mediterranean region.
Differences in physiographic conditions resulted in differences in the
development of landscapes and Cenozoic fauna between the western
Mediterranean — Pyrenean-Balkan, and the eastern Mediterranean — Balkan-
El'brus. Eveninthe Oligocene the southern part of the eastern Mediterranean
was under the influence of the central Asian continental Indricotherium
fauna. Later, the thermophilous faunal complexes persisted on the northern
and southern coasts of the western Mediterranean until the beginning of the
Pleistocene. These complexes consisted of primates, viverrids and
hippopotami. It has only recently been established that hippopotami and
primates lived in Transcaucasia in the Upper Pliocene- Lower Pleistocene.
At that time, the Azov Sea area was inhabited by some thermophilous forms
— elephant and antelope, but the complex, as a whole, was characteristic
of savannah and forest-steppe landscapes on the margins of temperate and
subtropical climatic belts.
The western Mediterranean in the Pleistocene was protected in the north
by the high Alps and the Carpathian Range, whereas the eastern sections
were exposed to masses of winter-cooled air from the Russian Plain.
The landscapes and fauna of Spain, Italy and Greece were not as strongly
affected by the glaciation of northern Europe as the landscape and fauna
of the Caucasus and the Crimea.
During the maximum, Dnieper stage of glaciation, a number of taiga
and tundra species of mammals extended their ranges over the Russian Plain
as far as the southern coast of the Crimea. These northern forms did not
460
463
464
migrate farther to the southeast, i.e., to the Caucasian Isthmus, because
the steppe zone of the Pleistocene on the Russian Plain made a sharp
northward turn in the northeastern Azov area. In addition, the opening of
the Manych strait during the Caspian transgressions (see Chapter II) also
influenced the distribution of the fauna.
The Mediterranean-type landscape is often identified by biogeographers
with the Recent landscapes of Spain, Sicily and Palestine, i.e., sun-burned,
rocky, dry slopes, grazed by goats, grown here and there with olive
groves, fig trees and orchards, supporting colonies of rabbits and stone
martens and occasional relict mouflons and wild goats. Actually in both
the eastern and western Mediterranean, relict landscapes of Tertiary
coniferous and broadleaf forests and mesophytic meadows of alpine- and
subalpine-type occur alongside xerophytic landscapes of garigue, phrygana
and upland steppes (see Il'inskii, 1937). Such characteristic species of
mammals as Asia Minor mouse, snow vole, Promethean vole, chamois,
Spanish, Alpine and Caucasian goats and macaca evolved and survived in
these mesophytic areas of the mountains. These species are probably as
old as piebald shrew, jerboa, gerbil, Selevinia and other animals of
the adjacent desert centers of speciation.
The highland landscapes of the Caucasus, like other mountain ranges
of the Alpine folded system, formed in the Miocene. Even if some
peneplanation subsequently occurred in the Pliocene, nevertheless some
of the ranges retained their prominence.
The Holocene complex of Caucasian mountain-forest mammals has,
therefore, many features in common with mountain-forest mammalian
complexes of the Alpine ranges of the western Mediterranean. In the main,
it evolved locally, rather than migrating as a unit, which accounts for its
high degree of specific and subspecific endemism. In the east this type of
fauna occurs in the forests of Talysh and on the southern slopes of the
Elburz. During the hot, dry stages of the Pliocene, it was affected by
southern faunal elements, and during the Pleistocene by influences from
the northeast, northwest and southwest. Of these, the faunal ties with
the south were the oldest, most extensive and most permanent.
The southwest Asian uplands are an independent center of evolution of
two subtypes of mammalian complexes of Pliocene age: the upland-steppe
and upland-desert complexes.
The differentiation within the southwest Asian complex of these two
subtypes is observable in almost all orders of mammals, and is related to
the sharply differentiated relief of the country and its climatic variability.
At the same time when biocenoses of the upland-desert type evolved in broad
intermontane valleys and narrow canyons, ecological assemblages of upland-
steppes and even meadow-steppes evolved on neighboring high plateaus and
ranges.
The influence of this southern complex on the Caucasian Isthmus was
undoubtedly exerted in several stages, the oldest being the Miocene-Pliocene
when the Hipparion fauna was dispersed over the Isthmus. It is probable
that considerable influence from the south continued through the Upper
Pliocene, but in the Pleistocene, particularly in the Upper Pleistocene,
migration from south to north was probably impeded.
In the north, the Caucasian Isthmus joins the southern Russian Plain
which, in the area of the Dnieper, Don and Volga watershed, has been dry
461
land since at least the Lower Miocene. The nucleus of the steppe flora and
fauna of this area formed in the Pliocene, according to pollen analysis
and paleofaunal data.
The early stages of settlement of the Ciscaucasian plains by immigrant
species from the Eastern European and northern Kazakhstan steppes might
have taken place in the Middle Pliocene. However, the large-scale
migration to the Caucasian Isthmus of mammals belonging to this steppe
complex undoubtedly occurred in the Pleistocene concomitantly with the
southward shift of landscape zones. During the humid climatic phases of
the Pleistocene, the mammalian complex of European and Asian broadleaf
and taiga forests could also have influenced the Caucasian fauna. These
forest mesophilous species could have dispersed along the valleys of the
Don and Volga, through the Balkans and along the northern coast of Asia
Minor.
The main features of the Transcaspian mammalian complex, i.e., the
central Asian desert complex, had probably developed by the Lower
Pliocene. The migration of representatives of the central Asian desert
mammalian complex to the Caucasian Isthmus could have come partly from
the northeast and southeast around the Caspian, and partly directly from
the east.
The Pleistocene and Holocene immigrants from the semideserts and
deserts of Turan to eastern Ciscaucasia — jerboa and gerbil — probably
appeared during the regressions of the ancient Caspian. The late, Holocene,
stage of immigration of a number of thermophilous southern mammals is
apparent in their ranges of distribution.
In our systematic analysis of Holocene mammalian fauna of the Caucasian
Isthmus we include, in addition to extant relict species, those species
which became extinct in the historical epoch under direct and indirect
anthropogenic influences. This approach gives a much more comprehensive
list of Caucasian animals than those compiled by other investigators.
As previously mentioned, the analysis of the Recent fauna was intended
to reveal two aspects of the distribution ranges: their origins and their
ages, which is to say, both the historico-genetic and ecological identity of
the species. Lacking paleontological data on some of the Caucasian species,
their ages were estimated on the basis of comparisons between the antiquity
and evolution of their preferred biotopes, on the one hand, and data on their
ecology and migrational ability, on the other.
The review of Recent and restored distribution ranges of Quaternary
mammals inhabiting the Caucasian Isthmus now or in the past (Maps 1-93)
and a comparison of these with paleogeographic data resulted in considerable
refinement of the scheme of genetic composition of the Caucasian Holocene
fauna given in Chapter III. In its complete form, the scheme is as follows:
PALEOGEOGRAPHY AND ECOLOGY OF CAUCASIAN MAMMALS
I. Tertiary (Pliocene) :
1) Caucasian, mesophilous, widely distributed;
2) Southwest Asian, xerophilous and thermophilous;
3) Eastern European, hydrophilous and steppe-type;
4)
465 Turanian, desert-type.
462
Ш. ЕР1е15юседе:
5) European, forest mesophilous;
6) European-Asian, steppe-type;
7) Turanian, semidesert- and desert-types.
III. Holocene:
8) South Asian, thermophilous;
9) Turanian, desert-type;
10) accidentally introduced.
The principal types of ranges of some indicator species of these complexes
are given in Figures 188-194.
The zoogeographic and chronological analysis of the Holocene fauna of
the Caucasus is given by orders.
INSECTIVORA
The Caucasian Isthmus is one of the richest areas in Insectivora species
in the U.S.S.R. We have identified 15 extant species from studies of
436 localities and 1,224 specimens (Maps 1-6). Modern compilations drawn
from the Caucasus and neighboring areas (Ognev, 1928; Bobrinskii,
Kuznetsov and Kuzyakin, 1944; Kuznetsov, 1950; Ellermann and Morrison-
Scott, 1951; and others) do not reveal occurrences of Brandt's hedgehog
of central Asia and Iran or of Diplomesodon or some white-toothed
shrews of the western Mediterranean in the Caucasian fauna.
The nucleus of the Caucasian Insectivora is composed of local mesophilous
species, such as the mole of the southern, Mediterranean type, Caucasian
and lesser shrew and gray white-toothed shrew. It also includes such widely
distributed species associated with mesophytic landscapes as European
hedgehog and water shrew, whose origin on the Isthmus is obscured.
Next in order of abundance is the group of species associated with the
Tertiary steppe and semidesert landscapes of the eastern Mediterranean,
e.g., the long-eared hedgehog, white-bellied white-toothed shrew and
pygmy white-toothed shrew.
Among occurrences of relict Tertiary species of the Russian Plain are those
of the Russian desmanwhich is limited to the northwesternmost part of the
Isthmus, the common shrew which is sparsely settled in the southeastern
part, and the little white-toothed shrew which is rare in eastern
Transcaucasia* and southwest Asia. These species migrated from the
north to the Caucasus in the Pleistocene.
Table 97 sets forth our proposed scheme of the genetic composition of
insectivores of the Caucasian Isthmus. **
* In Transcaucasia, this form includes Crocidura dinniki Ogn., whose systematic position is not clear.
** Space cannot be devoted to a criticism of the classification of shrews given by Bobrinskii, Kuznetsov,
Kuzyakin (1944) which was fully accepted by Ellermann and Morrison-Scott (1951), Peculiar
misidentifications have been made of the gray white-toothed shrew. Ognev (1928)erroneously identified
as gray white-toothed shrew ап alcohol-preserved specimen of long-tailed white- toothed shrew from northern
Iran (Mala-Kara), which is in the ZIN collections. Because of a semantic confusion (?!) Bobrinskii and
Kuzyakin misidentified the Pontian Crocidura lasia Thos. as the Far Eastern C. lasiura Dobs.—
an error which has resulted in a greatly confused nomenclature and has obscured discussions of the ranges
of these two species.
463
466 TABLE 97. Genetic composition of Caucasian Insectivora
Holocene
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Bae 2) ee | oa I> р o : o >
=f |< о] За < |= 2) Oi
мета | ae = - о ! -a] 8 Suds 3
a a oo =] Pt |= в та я > >
| оне | a |] a ‘a a ao) ag а 5
io ise <3 = 0 | oO oO ee ‚= т о
а мо | ее ees Se [А-В as)
Oia) ЕЕ Si alls te Mow’? [2] sg] = м |e
3 0 fas 5 an UO = a © = ны фФ =) Ия о
Gea 1 0 sila 5 = 5 о о зо о
OFljadnd|malh la & [5] Hola Ho <
Erinaceus europaeus L.
Hemiechinus auritus Gmel.
Talpa orientalis Ogn. ,...
Тъеацеа ао а ОА mance ort po amas
Резшапа Moschata sl ye ean
SHO ELE 2S alga lath of) FUR оо Оооо aye
SIL NS © a) ew levels sd seme our ie
5. Tadidiog "Sater getccuct casts, sbttioue-c
Neomys fodiems Schr. ......
Crocidura suaveolens Pall.
с. Тенсочет +Herni.w Seis. Fr
С. Tussula Herm: „вена sees
Са аа DOSS we зле Bens bane eons
Ca дара. a cae
Заре etruse Us Savy. 2%.
CHIROPTE RA
We have identified 27 Holocene Chiroptera species from 785 specimens
from 419 localities of the Caucasus (Maps 7-17). There are reasons to
believe that the fauna was somewhat poorer in the Upper Pleistocene because
of the southward retreat of the ranges of some thermophilous species caused
by the cold phases. These ranges were restored to their former areas in
the Holocene. It remains for future investigations to reveal new forms of
bats, so far unknown on the Caucasus because of the difficulties of bat-
collecting.
According to the compilations of Allen (1940), Tate (1942), Kuzyakin
(1950) and Ellermann and Morrison-Scott (1951), the subtropical families
Pteropidae, Rhinopomatidae, Embalonuridae, Nycteridae and Hipposiderinae,
464
which are characteristic of the southern parts of southwest Asia, do not
occur in the Caucasus. As a whole, however, the Caucasian bat fauna is
much richer than that of Europe and Asia. The ranges of most of these
Caucasian species belong in the Mediterranean region, sensu lato; viewed
ecologically, they can be divided into those of relatively mesophilous forms
and those of more xerophilous and thermophilous forms. The first occur
in areas of broadleaf forests where the bats often build their nests in hollows.
Their summer and winter habitats are mostly in marginal areas of the
mountain formations of the Isthmus; the latter forms prefer the landscapes
of low plateaus, foothills and hot lowlands.
TABLE 98. Genetic composition of Caucasian Chiroptera
Tertiary (Pliocene)
467
Holocene
Pleistocene
3 9)
[=] 3 5. 2
= Э > Е
= к > ie)
ея = = 1 He ue)
= 2 = oO el oO
ou |9 a v Sy |) ray |]! ef 3)
=) (©) oO ° a, a, = о = 3
= в |e = > ® ® ®
со. > “a uu = > an Е n 8
Species oa не || > ! О я о
Р те а || = = | э о | зо 5
ce (=! rp) = > 2 a
о > = 5 fey o 3 я o = Ф я
eS iia аа = & |= я Е! е|Е |=
ao) “Sala > [2 a Oo A : o РР >
г. < о o ! a5} mw 4} < ae |= ast —
|8 a, v ет фа i il
cso | 3 Qe я == = я о a а о =
A ча оно a os a| a an <| sd a я
x | || = 9 = 0 G,| о = oO ‘a © o
ва 2 a ls el eiaePilsiaw= | s
Sos su | 20 = O wm) 9 HD | ag о
Si | Oe | S & SB 15 8) 5 5S ¢ o| 5 4 9
Оз! ша H ja SJ) mo} a nl|H <
Rhinolophus ferrum equinum
SClimemtrcnct aus Nc coer or maces
Rune Мите ше у М атс eee ues
Rin, SWRVAIS Bl. с обооьовос в
в, bhp) роЗиеи@З Bechstary stun
Uns wile Si Reem Вы о об Gace 6c
Myotis bechsteinii Kuhl .....
М. emarginatus Geoffroy ......
Mls оо gmat ws MICE “Ge ea o se
Mi, MAY SRACMNOS KUM) 5 oS660 0c
о ПО i MIG wean im hs eee
Miniopterus schreibersii Kuhl
IE OL WE UN CEDIG. Ie oes oe o wo Ole
Barbastella darjelingensis Dobs.
B, DAT DSS Wlmg ЗМ 54 a ooo
NyCtailus Пола tial бое
INS GUS WI Me Paull, woeM tees Kets. обо ьо
ING MOC i Mlle Seite sg vavondhe selene alist
Vespertilio pipistrellus Schr.
Wo ме ами, 158593. Gis ВВ По 6 5 1
УЗО о SG ео ЗЕЕ
№. за ми Воларь еее oo Oe 6 aod
Е ее а acD
We Opis Rail) ИЯ S sae оо sls в
У
У
У
т
Оу ее АВА
ео ет ee Chee
порево ро во и. Пи
adarida taeniotis Raf. .....
465
468
The mesophilous group comprises most of the mouse-eared bats and
vespertilids, including the serotine vespertilio, which occurs more often
in beech forests, e.g., in Talysh, than on open plains.
The more xerophilous and thermophilous group consists primarily of the
horseshoe bats, barbastels and the Mediterranean pipistrel. The extent of
the influence exerted by European Chiroptera fauna is undetermined, but
it was probably not significant, no more so than that of the central Asian
desert fauna. Among the desert species, the distribution range of Ognev's
great bat is highly characteristic of the Tertiary hills in the Kura lowlands
of eastern Transcaucasia.
Only two species can be tentatively identified as Upper Pleistocene
immigrants from Europe: the giant noctule and the northern vespertilio.
Their appearance on the Caucasus at that time was possibly a second one
caused by the end of the glaciation.
The complete list, divided into genetic groups, is given in Table 98,
using the terminology of Ognev (1928) and Kuzyakin (1950).
Some species —Vespertilio bobrinskii Kuz. and У. ognevi
Bobr. — require additional study. The first should be compared with
V. (Eptesicus) nasutus Dobson, 1877, and the second with
V. (Eptesicus) sodalis Barr. Hamilton, 1910.
CARNIVORA
The Carnivora fauna of the Caucasus in the Holocene is comprised of
25 species; it was probably no richer in the Upper Pleistocene. Of these,
22 species survived until recent time. We have had available a wealth of
material for the zoogeographical analysis of this group (Maps 18-38):
extensive collections of fossil remains from 92 localities, skulls and skins
of extant species from 376 localities, observations and data from state fur
stations covering 257 localities, and 417 specimens. In comparison with
neighboring territories, the Caucasian Isthmus is rich in carnivores (Ognev,
1931, 1935; Bobrinskii, Kuznetsov and Kuzyakin, 1944; Ellermann and
Morrison-Scott, 1951). The absence of Viverridae from the Caucasian fauna
is noticeable, and is accounted for by the fact that the northern boundary of
their range lies somewhat to the south in Iraq. The family of honey badgers
(Mellivorinae) is also conspicuously missing, as they occur much farther
to the south and east. The rest of the Palaearctic carnivore families are
fairly well represented in the Caucasian fauna. The Caucasus lies outside
the ranges of some of the Canidae, e.g., some foxes of southwest Asia
(Vulpes ruppeli Schinz., V. cana Blanf., Fennecus zerda
Zimm.). True feline desert species (Felis margarita Loche and
F. caracal Schr.) do not occur in the Caucasus either. Most Caucasian
carnivores belong to ubiquitous forms of uncertain origins, which have,
nevertheless, been included by us in the group of Caucasian Mediterranean
forms because of the occurrences on the Isthmus of closely related forms
in the Middle and even in the Upper Pleistocene. These forms are wolf,
fox, European brown bear, badger, and lynx.
466
(469) TABLE 99. Genetic composition of Caucasian Carnivora
Holocene
3 3 o
э 3 Э В. Е
— о — —
= ae) = a о
nan = St „= 1 rt ~~
=) 2. a с Фо = 1 ro)
ov [<] a. o [2] (ay ||: a, > Oo
= 0) oO (©) 5. я Oy = о FA 5
РЕ С о a
С ey | ca] = = Е |“ Я ® |2)
Species о > 8 но > > ~ |U wv о Gn 8
a Е oO oO + 3 =]
ov — ® n = =) os) oO я
a MB ‘+ - с [$ & с. — |= о, “4
Я + на Е > А = Gs)
и м || = G Gh ep || =
> о oO т
|| а 9 с > < ee Е м
зо a oe |= = = в а И >
a ORK] ea aA a ® Е
a VU [$ 5 о < is) < go a < Чем (9
Sas 2 4 Ее || Ф = o Е ® ®
oF |Ез а | & a. яз a a 9 |] 34
Е ЕЕ ile
Oa очшз & wm m |H @ [22] Ed <
Grats rans ми.
CHUMP ANS Ley Gh ete tale a. hee оровооо =
Varlipesrv wipes и и © .s adele =
Wol CORB ENCE NST lo. cache оо. В ood
Нузепайпуаена т: еее
ПОРТ (956) И ways о оО Во о
Маре ищаыье: Ти. осо чан
п MOHTAL LET pg co ei lecelieLiclbe! aula cinc
Vormela peregusna Guild. ...
PP MEO РОМ jh 25 6 6 ulls
P. eversmanni Less. ..... dio od
В ОИ ee о alo Giallo
INE SCY NCUA Тре о оса
да) лета ие к рвы еее
“ Рашедера 515019 Ibe enone os
SPV eOLy, ana mtn tat: о
о О оао
СОЭ UVC CR ооо роовов
Me Энен бат ов once a
р. CMAs Gill, . 526455 ее
По ВВС рос
Otocolobus manul Pall.
* Acinonyx jubatus Schr,
Note. ?—species whose occurrence on the Isthmus has not been definitely established; * —species which
became extinct in modern time.
The southern (southwest Asian-Mediterranean) species of steppe and
desert landscapes are represented by pine marten, tiger polecat, African
wildcat and cheetah).
The Pliocene eastern species (Turanian-Afghan) are represented by
the manul.
The Pleistocene European immigrants from the north and southwest
are easily recognized by the poor development of their ranges, which
wedge out toward the southeast. These forms include pine marten, mink,
European wildcat and, possibly, the ''European-type'' bear of the Greater
Caucasus.
467
АП the immigrant carnivores of the Holocene originated in the south and
are relatively thermophilous. Their late appearance on the Isthmus is
confirmed by modern ecological data, the development of their ranges and
the absence of their remains in Pleistocene bituminous deposits on the
Apsheron and in the caves on the upper Rion. The jackal, striped hyena,
tiger, lion and jungle cat belong to this group; their origin on the Caucasus
has been discussed in Chapter III.
Most of the nomenclature employed for the list of carnivores given in
Table 99 follows the usage of Russian authors.
469 LAGOMORPHA
Two species of Lagomorpha emerged on the Caucasus in the Holocene —
the European hare and the Armenian pika. Indications from Recent
ecological data, from the characteristics of their ranges and from Cenozoic
fossil hares and pikas, found on the southern Russian Plain and in
Ciscaucasia, are that they were ancient Pliocene (Miocene) immigrants to
the Isthmus, which evolved in the eastern Mediterranean xerothermic
landscapes of the Tertiary. Of the more southern forms, the Arabian hare
(Lepus arabicus ЕЪг. ) does not occur on the Caucasus, and of the
western Mediterranean forms, the rabbit (Oryctolagus cuniculus L.)
is absent.
The Pleistocene migration of the blue hare and small pika from the
north to the Caucasus has not been proven.
The two Holocene Caucasian species are Tertiary forms of eastern
Mediterranean origin and their Recent ecology indicates their relatively
xerophilous character (Table 100, p.471).
470 RODENTIA
The Holocene fauna of the Caucasus includes no less than 50 species of
rodents. The specific composition of the Caucasian rodent fauna can be
regarded as completely known from the 10,623 specimens which we studied
and from data from 1,770 collecting localities of extant forms. The Middle
and Upper Pleistocene rodent fauna was probably not much richer than the
postglacial fauna.
The variation of the zoogeographical and ecological composition of the
rodent fauna and its considerable geological antiquity permit a number of
reliable conclusions on the particular features and developmental patterns
of this faunal group. Taking stock of the faunal complexes of neighboring
territories and their potential influence on the Caucasian fauna, it becomes
apparent that a number of rodent species do not occur on the Isthmus.
This review and comparative study of rodent fauna are based on works by
Ognev (1940, 1947, 1948, 1950), Bobrinskii, Kuznetsov and Kuzyakin
(1944), and Ellermann and Morrison-Scott (1951).
468
471
The following species are missing from the Caucasian fauna: from the
Eastern European steppe complex — marmot*, gray, big or red-cheeked
suslik and Eversmann's hamster; from the European forest and mountain-
forest complexes — common squirrel, dormice of the genera Eliomys
and Muscardinus, the Balkan Dolomys, common vole, root vole and
a number of other subterranean voles of the genus Pitymys; from the
central Asian desert and southwest Asian upland-desert complexes — large-
toothed suslik, long-clawed suslik or ground squirrel, jerboas of the
genera Pygeretmus and Jaculus, mouse-like dormouse and
Betpakdala dormouse, many species of gerbils of the genera Gerbillus,
Psammomys. Tatera and subterranean rats of the genus Nesokia.
There are also some western Mediterranean species that do not occur
on the Caucasus: African porcupine (Hystrix cristata L.), mouse
(Lemniscomys barbarus L.), Arvicanthis niloticus Desm.,
Acomys cahirinus Desm., and other species.
Upper Tertiary (Pliocene) mesophilous species of the humid broadleaf-
forest and mountain-meadow landscapes of the eastern Mediterranean
comprise the largest group within the Caucasian rodent fauna:
Persian (Caucasian) squirrel Broad-toothed field mouse (Asia Minor mouse)
Fat dormouse Promethean vole
Forest dormouse Water vole
Black rat Caucasian snow vole
House mouse Long-tailed snow vole
Caucasian yellow-spotted mouse Pine vole
Common field mouse
The species referred to here as house mouse includes a number of wild
forms, some associated with biotopes of the mountain, foothills and
lowland-forest zones: Mus musculus formosovi Hept. and
M. musculus abbotti Wat.; some associated with biotopes of the
steppe and semidesert: М. musculus hortulanus Nordm. and
M. musculus tataricus Sat. Those large Caucasian mice of the
Apodemus fulvipectus Ogn. type (A.flavicollis ponticus
Svirid., A.f.saturatus Neuh., A.f.argiropuli Vin. etArg.)which
inhabit the forests of the Caucasus, and which grade into a smaller, lighter-
colored form with a small spot on the neck which inhabits the semidesert
zone of eastern and southern Transcaucasia (i.e., the A. arianus Blanf.
type), are regardedas a polymorphic species of the eastern Mediterranean
(Caucasian) yellow-spotted mouse.
The ancient xerophilous species of eastern Mediterranean origin include
the following:
Asia Minor suslik Asia Minor gerbil
William's jerboa Vinogradov's gerbil
Asia Minor hamster Transcaucasian mole vole
Migratory hamster : Asia Minor snow vole
Mouse-like hamster Lesser mole rat
Red-tailed Libyan gerbil Porcupine
Persian gerbil
* Tt is possible that the bobak marmot inhabited the Ciscaucasian steppes in historical time.
469
472
The time of the emergence of these ancient species in different parts of
the Isthmus and the characteristics of their dispersion vary, as the examples
given in Chapter III show. :
The European beaver and Ciscaucasian mole rat are representative
of Eastern European Tertiary species, hydrophilous in nature and adapted
to steppe ecology. Without pursuing the question of the origin of European
beaver, it is assumed that the species inhabited the Caucasian Isthmus and
Russian Plain as early as Pliocene time.
It is more difficult to place the origin and emergence of mole rat in
Ciscaucasia. In the absence of fossil mole rats on the Isthmus and any
proof of their migration from Asia Minor, they must be considered
descendants of the Miocene-Pliocene mole rats which inhabited the southern
Russian Plain. Immigration of the Russian mole rat and the giant Russian
mole rat to the Isthmus could have taken place at the time of deposition of
the Balakhan beds, i.e., in the Middle Pliocene, and they could have
survived the subsequent marine transgressions in the Caucasian foothills.
The sole Caucasian representative of the ancient Turanian and Afghanistan
desert species is the small five-toed jerboa. Its distribution range, which
includes eastern Ciscaucasia and eastern and southern Transcaucasia, is
an interesting example of the influence of the ''Aral-Caspian fauna."
A fairly large group of Pleistocene immigrants of various ages and
origins canbe identified from a comparison of the ranges and
palaeogeographic data.
There are at least seven generally mesophilous, ''European"' forest
species:
Northern birch mouse Common hamster
Harvest mouse Common vole*
Striped field mouse Common red-backed vole
Yellow-necked field mouse
The ranges of these species on the Isthmus are rather poorly developed.
Where they penetrated deep into Transcaucasia, the ecological channels
of their dispersion were either the floodplain river marshes of the piedmont
valleys or the mesophytic meadows and coniferous forests of the uplands.
Their routes followed the Black and Caspian sea coasts, bypassing the
Caucasian Range on the east and on the west. Examples are the harvest
mouse, striped field mouse and common hamster. The range of the
European yellow-necked field mouse of the Apodemus flavicollis
samariensis type is confined to the tugai forests of the lower Terek and
Sulak in eastern Ciscaucasia. Indeed a puzzling phenomenon of distribution.
Since it is doubtful that the species is endemic, it would seem that
it could only have migrated to this region from the lower Volga. The most
probable time would have been the Upper Pleistocene; the desert landscapes
which formed in the northern Caspian area in later, postglacial, time could
not have carried the migration of this species.
Immigration of three eastern xerophilous species of the southern
Kazakhstan semidesert and of the central Asian desert complexes occurred
in the Pleistocene: thick-tailed three-toed jerboa, northern three-toed
jerboa and little earth hare.
* Bones of this vole which have recently been found in Acheulean beds of Kudaro I cave indicate that the
occurrence of this species on the Caucasus dates from ancient time.
470
(473) TABLE 100. Genetic composition of Caucasian Lagamorpha and Rodentia
Tertiary (Pliocene) Pleistocene Holocene
3
= n
о =) =}
= ig) о
[22 ao} a =
= |5 a. о 0 = а
Speci оо 2 = я [=] =
Becies FN dee, 5 = a Е | 3
= a . 3s] = и ® 5 ®
aele боя 2 Sie |S 1s &
a о 3 ява. | = я ©.
ааа [5 cr op) о | os
xc a] 3 Es} uw 2 ojo” = о Ae!
> о|9 0 5|<%8 1 В и
вы Ее ее а ао а ща
ов | в ae La sia & a ®
пана < (23
SS ge Ma 5 = © 5 a oe) = Е
ее | se
Odtg|na|[m a = шв ша езч|о = <
Lagomorpha
Lepus europaeus Pall. ......
ООО Bs ооо обаюо
Rodentia
Sciurus anomalus Guld. ne
Citellus pygmaeus Pall. ....
Ce оО и ie once creas
Ea Gaston Роем
GUIS BIBI зоб 564000 cco
Dyromys nitedula Pall. ....
ето ва мера Wy aie cee
SHpbietwlainiawPall a: Bote Siem be
SHICAINIC ais Car Ур eee lakes
Allactaga elater Licht. .....
даем Ра. ое co ketene
АИ ато Ве
Alactagulus acontion Pall...
Scirtopoda telum Licht. ....
DAPwUS Зато Pell, ob oncag oc
Spalax microphthalmus Са.
5. gigantews Nehry н.о,
S. leucodon Nordm. ....... BP
RB bG USM Ratt Wise Los fuses his qs oe
В. norvegicus Berken...... са
MEG о 6 оо бобов
Micromys minutus Pall.
Apodemus agrarius Pall. ....
ИЗ OPT, orden ou оо
A. flavicollis Melch. ...... |
а ВУИ WS) Vg Бор бобов
А. mystacinus Danf. et Alst. ..
Mesocricetus auratus Water.
(Сони ООС ооо
Cricetus migratorius Pall. .,
Calomyscus bailwardi Thos,
* Rhombomys opimus ......
Meriones meridianus Pall,
MivenythrowrusiGrayeyt). мя.
М; решен Вал ve, ды ances
М. всататт ее Pall secs «
Note. Asterisk indicates species which became extinct in historical time.
471
Accidentally introduced
TABLE 100 (continued)
Holocene
Species
Southwest Asian, xerophilous
European - Asian, steppe-type
and thermophilous
East European, hydrophilous
Caucasian, mesophilous
and widely distributed
and steppe-type
Turanian, desert-type
Turanian, semidesert-
South Asian, thermophilous
Accidentally introduced
European, forest
mesophilous
and desert-types
Turanian, semidesert-
and desert-types
Meubibaie@kTe mie Бо: ее m tia
M. vinogradovi Hept. ......
NEE TOMES SPe cw о мо беоне
Lagurus lagurus Pall. ......
Ellobius talpinus Pall. .....
EMMne Se. enismkossy te-a.tanske tens &
Prometheomys schaposchni-
KOMP Sat А ОЕ sear ae
(474)arvicola terrestris L. .....
Microtus nivalis) Маг... .....
НЕ Пе ое c
ODE ыы По оном eae fee
Majorim Bhost’ .... F365 5.2
Arvid l US EAUeee.encneear cheat, @
ое ta ира Peewee es cee -
Clethrionomys glareolus Schr.
Hiystrix leweuna Sykesp: Shr.
ae EP:
Their Pleistocene age is evident from the Upper Pleistocene occurrence
of their remains on the lower Don, indicating that these species migrated
from the Transcaspian deserts far to the west no later than the time of the
post-Khazar sea regression. Partial survival of the Khvalynsk transgression
was possible on the southern shores of the Terek bay. Again in the Holocene
there was another wave of immigration of these species and of Turanian
desert species to the Terek-Kuma semidesert and the Manych area. The
great, midday and tamarisk gerbils are representative of these Holocene
immigrants from the central Asian deserts, as shown by the absence of
their remains from Quaternary alluvial beds of the lower Don and by the
sparseness of their Recent distribution. The great gerbil became extinct
in Ciscaucasia in historical time.
One would expect that new southern rodents would have emerged during
the xerothermal postglacial epoch in the southeastern part of the Isthmus,
in addition to this second wave of immigration of desert species to the
northeast. However, no such instances are known with the possible
exception of the distribution of Asia Minor gerbil on the Kura-Araks lowlands
and the local dispersion of Persian gerbil, steppe vole and some other
species caused by anthropogenic changes in the landscape.
472
474
475
The Norway rat is the only accidentally introduced species which has
inhabited the Isthmus in historical time; this species and black rat are
widely distributed over railroad, water and air routes. The penetration
of house mouse of the northern type into the Caucasus continues.
The complete genetic composition of the rodent fauna is given in
Table 100, employing the basic nomenclature accepted by Russian authors.
Thus, the great jerboais referred to as Allactaga jaculus Pallas,
and not as А. major Kerr.; the thick-tailed three-toed jerboa* as
Alactagulus acontion Pallas, andnot as A. pumilio Kerr.; the
red-tailed Libyan gerbilas Meriones erythrourus Gray, and not
as M. lybicus Licht.; the porcupine as Hystrix leucura Sykes,
and not as Hystrix indica Kerr., etc.
PR lows ODA Cl YIEvA
According to collection material and data drawn from ancient historical
sources, two or three species of horse lived in the Caucasus in postglacial
time. The origin of tarpan remains obscured by unanswered questions on
the morphology and evolution of horses of the Old World. Adopting the
generally accepted view, we have included it in the group of Pleistocene
immigrants from the northern steppes. The kulan is considered
a Holocene immigrant from the south. It seems likely that
postglacial evolution of horses proceeded differently in the north and in the
south. Ciscaucasia was inhabited by horses of the Eastern Sur one ah plas, feyu
whereas Transcaucasia was inhabited by southwest Asian horses. } Е
As is the case with some rodents and carnivores ‚ perissodactyls
occurring in different parts of the Caucasus had different origins. Examples
are the kulan and the southwest Asian horse, whichinsouthern Transcaucasia
must be regarded as local Tertiary species, but which ineastern Transcaucasia
and Ciscaucasia must be regarded as Holocene immigrants.
Table 101 shows the different positions occupied by Caucasian
perissodactyls in our scheme.
AR DIODACT Y LA
The Caucasian Isthmus occupies first place inthe U.S.S.R. in the
number of artiodactyl species and individuals per unit area. It is a large
center of speciation and development of this order.
There has been no migration of the arctic forms, reindeer and musk ox,
to the Caucasus. And the ranges of southern thermophilous species (such
as Asia Minor and Mesopotamian fallow deer belonging to the genus Dama)
and of a number of southwest Asian gazelle species (Gazella dorcasL.,
С. gazella Pall., С. leptoceros Cuv.) do not extend to the Caucasus
in Recent time.
Of the mountain forms of central Asia, no markhor [Capra falconeri
Wagner] occurs in the Caucasian fauna; and of the Western European forms
no Ammotmapus lemrvna Pall | Capralisinaitica Ehr,,
C. ibex L., or C. pyrenaica Schinz. are found either.
* [This is the correct translation for the Russian "emuranchik," but the author apparently meant "little
earth hare" ("zemlyanoi zaichik") ... see Table 103. ]
473
TABLE 101. Genetic composition of Caucasian Perissodactyla and Artiodactyla
_ Tertiary (Pliocene) Holocene
2 o
5 2 с. n
8 5 > => 3 ae)
= о > |= о =
a ие = 1 a = 5 =
5 a, = © = o
оэ|о a. Ф 5. ' 1
ao] [2] а. a = 5 a =.
5 3 И = ES К o Е o =
— nn
pecies = = 3 > 1 7 о — В Е
с Е ec ial pee) a4 — - Uv Vv uo) e
o =P = vo oO a = = а „= =
2 Ss ta - с. a oO a Е - Е =)
Sst | = ich Ss р Be
ao] 2. |9 = o OF tes a o - vo >—
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>. о o = < —
те |5 Ф - ‚ Oo ' г. З . =
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о < = = шо Е 5 о о. = = = Е
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Perissodactyla
* Equus hemionus Pall.......
* Е. caballus gihelini АВЕ
ЗЕ cabia ии 0.
Artiodactyla
SUS ASC HOP ag нА. > кл
Cervus elaphus L.
GP ULGES JANCIS Alas а «ap aa
Gapreolus capreolus Lo... .
Seal Wiel раза Fara En) Wares eons 6 tie
Gazella subgutturosa Guld....
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Note. Asterisk indicates species which became extinct in historical time.
There was a gradual impoverishment in Artiodactyla on the Isthmus
in the Quaternary, in spite of immigration of new species from without.
Fifteen to sixteen species existed there in the Pleistocene, but by the
Holocene, there were only thirteen, of which only eleven survived into
recent time. Most of the Caucasian artiodactyls are local mountain-forest,
often endemic, forms. The endemic forms include the Caucasian chamois
and the east Caucasian and the west Caucasian goat. The group of widely
distributed, generally somewhat mesophilous forest species is comprised
of boar, red deer, roe deer, bison and primitive bull. It is possible that
elk should also be included in this group, once its southern origin has been
established.
474
The southwest Asian xerophilous species, which are mainly associated
with upland-steppe landscapes, are represented by Bezoar goat and Asia
Minor mouflon.
476 The Pleistocene immigrant group includes the northern-type elk, which
may have made a second appearance in the forests of the Caucasus, and
the saiga. The latter species reached the maximum development of its
distribution on the Isthmus in the Middle Pleistocene. This group should
probably also include the large form of roe deer which occurs in
Ciscaucasia.
The goitered gazelle is the only late — postglacial — immigrant; it
penetrated eastern Transcaucasia from the south, and eastern Ciscaucasia
possibly from the east.
The genetic composition of Holocene artiodactyls of the Caucasus is
given in Table 101.
The results of this paleogeographic and zoogeographic analysis of the
Holocene mammalian fauna of the Caucasus can be summarized as follows:
1. The Holocene mammalian fauna of the Caucasian Isthmus consisted
of 136 species of Insectivora, Chiroptera, Carnivora, Lagomorpha,
Rodentia, Perissodactyla and Artiodactyla, of which 124 species survived
into recent time.
2. The nucleus of the fauna — up to 36 % of the species — consists of
local (predominantly mesophilous) and widely distributed forms of Pliocene
age, They subsequently evolved on the Caucasian Isthmus and in adjacent
areas of the eastern Mediterranean where the landscape retained its original
(mesophytic) character because of proximity to the sea, warm climate and
a particular relief.
TABLE 102. Genetic composition (number of species by type of origin) of the Recent (Holocene)
mammalian fauna of the Caucasus, by orders
Tertiary (Pliocene)
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481
3. The next largest group — up to 27% of the species — is that of
southwest Asian xerophilous immigrants of Tertiary age.
4. Eastern European steppe and hydrophilous forms, some of which
migrated to the Caucasus in the Pliocene and some in the Pleistocene,
account for 12.5% of the species.
5. European mesophilous and forest species, which appeared on the
Caucasus mainly in the Pleistocene, constitute up to 11% of the species.
6. Postglacial immigrants, consisting of thermophilous species which
could not live under conditions of deep snow cover, and which migrated from
the east and the south (central and southwest Asia), comprise up to 7%
of the species.
7. Accidentally introduced species which appeared in historical time
amount to less than 1%.
All of these data, which differ considerably from the opinion of other
authors, have been summarized in Table 102. (р. 475).
The proposed scheme of geographical zonation of the mammals of the
Caucasian Isthmus is based on the results of the zoogeographical mapping
and the analysis of the mammalian fauna given in this chapter and Chapter III.
The generalized contours of the ranges of indicator mammalian
complexes, both those that are relatively stenotopic to the Caucasus and
those that were introduced, were employed to delineate districts,
subdistricts and sectors (Figures 188-194). In preparing the zonation, the
paleogeographical significance of territories and species was considered
first, rather than the simple numerical preponderance of animals in one
or another zoogeographic group. Thus, for example, the Caucasian
mountain-forest district includes, in addition to the Greater and Lesser
Caucasus, the eastern slopes of Talysh and the northern slopes of Elburz,
even though only four clearly autochthonous species of the ancient
mammalian complex of the Caucasus remain in the latter two sectors.
The rest are Pleistocene and Holocene forms. The entire steppe area
and part of the forest-steppes of Ciscaucasia belong to the Eastern European
steppe subregion of Pleistocene immigrant fauna, inasmuch as no
autochthonous Caucasian elements of Pliocene age remain in the Holocene
fauna of this area. Our subdivision of the territory is not based on as
many units as the zonation schemes of other zoogeographers, who based
their schemes more on geomorphology and geobotany, rather than on the
zoological material. Nevertheless, the boundaries of the phytolandscapes
figure significantly in our scheme because of the importance of the
relationships between index species of mammals and the phytolandscapes.
In view of the fact that the greater part of the Caucasian Isthmus belongs
in the eastern half of the Mediterranean region of the Palaearctic (i.e., the
eastern Mediterranean subregion), we include the northern part of the
Ciscaucasian plains in the Ciscaucasian subdistrict of the subregion of
Eastern European and northern Kazakhstan steppes, and the northwestern
corner of eastern Ciscaucasia in the Terek-Kuma sector of the central
Asian desert subregion. The entire Greater Caucasus, the northern slopes
of the Lesser Caucasus and the eastern slopes of Talysh are included in
the Caucasian district, which is further divided into the Asterabad
or Talysh- El'brus andthe Dagestan subdistricts. The Lesser
Caucasus uplands (the Armenian Highland) is included in the Sevan
subdistrict, and the semideserts and steppes of eastern Transcaucasia
483
are included in the Kura subdistrict of the southwest Asian district of the
same eastern Mediterranean subregion. The Talysh upland-steppe and the
middle Araks valley are regarded as the main sectors of the southwest
Asian district. The general scheme, the historico-geological characteristics
of the districts, subdistricts and sectors and of their mammalian complexes,
are given below and summarized in Table 103 (p.500) and Figure 195 (p. 515).
EASTERN MEDITERRANEAN SUBREGION
CAUCASIAN DISTRICT
(District of mole and pine vole)
The definition of the Caucasian district is based on the distribution
ranges of some index species of the Caucasian mammalian complex. These
ranges almost entirely cover the mountain forests, upland meadows and
rocky passes, the wooded parts of the inclined piedmont plains of
Ciscaucasia, and the lowland forests of Transcaucasia. The northern
boundary of the district is in the area of the inclined piedmont plains.
In the Ciscaucasian steppes there are two isolated sectors which belong
to this district — Stavropol and Pyatigor'e (Figure 195). In eastern
Transcaucasia the southern boundary of the district runs along the latitude
of Shemakha and Ivanovka in the Agri-Chai and Alazan valleys and near the
Kakhetian Range at Telavi. In Kartalinia and South Ossetia, the boundary
is on the northern edge of the dry Gori depression. It passes at the foot
of the Surami Range where it turns sharply east to the foot of the Trialet
Range, continues above Tbilisi and on through the lower Khram valley at
the foot of the Shakh-Dag, south of Shamkhor and Kirovabad. In the
southeast, the boundary bypasses Karabakh at altitudes of 600-700 m near
Martuni, and encompasses Karyagino and part of the Okhchi-Chai basin.
It takes a turn to the north and northwest, passing east and northeast of
the high steppes and meadow-steppes of the Armenian (Sevan) Highland.
The southern boundary of the Lesser Caucasus subdistrict of the Caucasian
district follows the divides of the Shakh-Dag and Pambak ranges ina
482 westerly direction, passing somewhat south of Kirovakan, and, skirting
northern Dzhavakhetia, enters eastern Turkey along the divide of the
Adzhar-Imeretian ridge. Its western boundary extends from Trabzon
to Sukhumi along the Black Sea coast; shifting slightly to the east, it
coincides with the boundary of the Ciscaucasian subdistrict of immigrant
steppe species.
The Caucasian district includes the isolated Asterabad Talysh-El'brus
sector which lies in the far southeastern part of Transcaucasia. The
boundaries of this sector almost completely coincide with the contours of
the forest zone on the eastern slopes of Talysh and the northern slopes of
Elburz.
The main massif of the Caucasian district encompasses the Dagestan
subdistrict, located between the mesophytic landscapes of the marginal
ridges of the eastern Caucasus (Figure 195).
484
483
It should be noted that where the boundary of the Caucasian district
passes at the foot of ranges and foothill plains certain parts of it comprise
a broad, transitional zone where animals of the steppe plains and of the
mountain-forest landscapes coexist sympatrically. Contrary to Kuznetsov's
opinion (1950), this zone, even in Ciscaucasia, has little in common with
the forest-steppe of the Russian Plain.
Most of the district has been dry land at least since the Oligocene,
gradually increasing in size on the periphery. Large mountain ranges
formed in the Miocene. The central parts of the Greater Caucasus mountain
system are composed of Jurassic slates and volcanic rock. The diluvium
and soil cover are well developed only on the high plateaus; in the canyons
the bedrock is exposed as rocky cliffs and talus. The marginal ranges are
composed mainly of sandstones and limestones, which in the foothills are
overlain by deposits of the Cenozoic seas and alluvial-diluvial boulder trains.
The Lesser Caucasus and Talysh present similar geologic phenomena
but calcareous deposits occur more rarely in these areas. As the
orogenesis of the Pliocene and post-Pliocene waned, stages of peneplanation
of the ranges alternated with mountain-building and glacial cycles.
The contemporary types of vegetation in the Caucasian district date
from the Lower Pliocene at least. The Recent zonation of climate and
altitudinal vegetation zones were formed by the first half of the Quaternary.
The contemporary landscapes of the Caucasian district which were
formed in the Holocene are not homogeneous over its vast area. In the
west, the Trans-Kuban piedmont plain is somewhat hilly in places and is
dissected by numerous, almost completely non-meandering streams.
The upper half of this plain is covered by oak and smoothleaf-elm
forests alternating with wide sections of steppe meadows. This type of
landscape does not occur in central Ciscaucasia; it reappears on the Terek-
Sunzha inclined plain, and finally disappears in the east near Sulak, east
of Gudermes.
In Transcaucasia — Colchis, Alazan-Agrichai, Khachmas and the
Lenkoran piedmont lowlands — lowland forests of a special type have
developed in which oak, Caucasian wingnut and smoothleaf elm covered
with thorny liana predominate. The marshy areas are characterized by
European alder. The abundance of nut shrubs and trees on these lowlands
favors the existence of rodents, ungulates and carnivores.
The slopes of the Chernye Gory, the northernmost ridge of the Greater
Caucasus, are covered with humid beech—hornbeam forests. Relict grassy
steppe vegetation covers some of the calcareous rock outcrops.
The next ridge, the Skalistyi, is characterized by subalpine meadows
in mountain passes and by mountain pine forests which replace the beech
forests on the southern slopes. Sections of mountain xerophytic vegetation
occur in the valley bottoms of the Kuban, Teberda and other rivers,
particularly in the longitudinal-latitudinal valleys between the El'brus and
central Dagestan.
Most of the mountain slopes of the lower and middle belts of the southern
slope of the Greater Caucasus, of the northern slopes of the Lesser
Caucasus and of eastern Talysh are covered with beech and hornbeam
forests, which provide seasonal food for forest rodents and ungulates.
485
484
Fir and pine forests are developed in the upper mountain-forest Бей of
the western Caucasus, and fir—maple forests on the southern slopes.
The well-defined rocky relief is a favorable habitat for snow vole and
chamois, which are plentiful in the area. As arule, the upper edge of the
forests is characterized by high-mountain oak and birch. Caucasian
rhododendron which wedges into the upland-meadow zone is also typical.
The alpine zone proper is completely developed in the western and
central parts of the Greater Caucasus and on the highest ridges of the
Lesser Caucasus with mesophytic meadows, taluses and rocky passes,
and glaciers at altitudes of 2,800-3,500 m. In the uplands of Talysh the
upper edge of the forest lies on the boundary of upland-steppe and upland-
xerophytic vegetation formations. In the eastern part of the Greater
Caucasus the subalpine meadows usually grade into areas of dense
bunchgrass formations, particularly sheep's fescue and matgrass, which
are typical for eastern alpine meadow formations. The steppe development
is even more pronounced on the median-altitude plateaus of Dagestan where
low precipitation and physiological dryness of the thin soil layer prevail,
and where upland-steppe meadows and meadow-steppes with low grasses
became the predominant formations. The turf-covered sections and slopes
of the plateau, which have a plentiful food supply of grasses and a rapid
run-off of rain- and snow-water, are particularly favorable environments
for subterranean rodents such as hamster and vole. The main food
resources of highland ungulates are also located in this zone.
The Recent mammalian complex of the Caucasian district mainly consists
of local Tertiary mesophilous forms which are associated with highland
meadows and mountain and lowland forests. This mammalian complex and
the ranges of its components developed out of local speciation and biocenotic-
forming processes which occurred in the Neogene and the Quaternary.
Immigration of animals from adjacent territories to this district, mainly
to the piedmont transitional belt, took place during the Quaternary.
The Holocene mammalian complex of the Caucasian district consists of
88 species, most of which belong to genetic group 1 (see p.462). There is
little representation from group 2, but somewhat more from group 3.
No Turanian forms, i.e., representatives of group 4, appear in the complex.
The nucleus of group 5 is fairly strongly developed, but group 7 is
completely unrepresented. There are a few occurrences of species from
groups 8 and 10(Table 103, p.500).
The index species are local representatives of group 1°
mesophilous Pliocene species comprising the Caucasian faunal nucleus
and the widely distributed forms, totalling at least 48 species and accounting
for more than one half of the total number of animals of the district
(Table 103).
Seem from the genetic and ecological points of view, this is not a
homogeneous group. It includes animals associated with the mesophytic
forests and meadows, which are fairly widely distributed in the broadleaf
forests of Europe, such as lesser shrew, fat dormouse, roe deer and red
deer. The list of endemic Caucasian highland species associated with alpine
mesophytic meadows is more specific. Caucasian birch mouse, Promethean
vole, long-tailed snow vole and Caucasian snow vole are characterized by
their mountainous ranges. In addition to these rodents, the larger, more
dynamic, but still stenotopic, species which are associated with mountain
486
485
relief, rocks, alpine meadows and mountain forests (Caucasian goat and
chamois) are also indicative of the Caucasian complex of mammals.
In the Stavropol and Pyatigor'e sectors and the Dagestan and Asterabad
subdistricts, the species of group 1 are poorly represented and have not
permeated the complex, mainly because of anthropogenic influences and
postglacial changes in climate and landscape. As examples, there are no
deer, goat, chamois, Promethean vole or long-tailed snow vole in the
Stavropol, Pyatigor'e and Talysh sectors; Promethean vole does not occur
in the Dagestan sector, etc.
Among the species in this group, European beaver and Caucasian
mountain marmot recently became extinct, but the black rat has survived.
Xerophilous and thermophilous Pliocene species (group 2) are represented
by Chiroptera, Carnivora, Rodentia and Ungulata which migrated to certain
landscape areas.
Since a number of species in this group could cross the ecological
barriers on the periphery of the mountain system, these species could
also inhabit some areas in the interior of the mountain system, such as the
longitudinal valleys of Svanetia, Kabarda and Digoria.
There are at least 16 extant species, of which the most common are
horseshoe bat, pine marten, European hare, Dagestan hamster, porcupine
and Bezoar goat.
The group of forest and mesophilous forms is comprised of Pleistocene
immigrants of northern (European) origin (group 5). For the most part their
occurrences are confined to the zones of mountain forest and highland
meadows of the Greater Caucasus. The habitat of the common shrew is an
example. Other species are more widely distributed, e.g., the Caucasian
[subspecies of the European] wildcat and pine marten, which, however,
do not occur in central Dagestan and Talysh. The pine marten is also
absent from the eastern part of the Lesser Caucasus. Ciscaucasian
subspecies of mink and field mouse occur in the district only in the foothills
of the northwestern Caucasus. The distribution of the common vole and
the Pontian common red-backed vole is somewhat peculiar: the range of
the first is primarily connected with the highland meadows, but does not
extend to western Transcaucasia; the partial range of the latter extends
from Asia Minor to the northern slopes of the Adzhar-Imeretian ridge.
The distribution of Caucasian elk, which became extinct in the last century,
was probably limited in the Pleistocene to Ciscaucasia and western
Transcaucasia.
It is exceptional that the little suslik, a representative species of group 6,
should occur in this district; it inhabits the dry longitudinal valleys of
the northern slopes of the Bolshoi Range along with Caucasian snow vole
and Caucasian goat.
The occurrences of Recent southern thermophilous immigrants (group 8)
— jackal, tiger and hyena which inhabit the periphery of the district —
are noteworthy.
The sole species from group 10, the Norway rat, inhabits the lower
mountain-forest belt of Ciscaucasia and the lowland forests of eastern and
western Transcaucasia. The species is particularly numerous in the
marshes of Colchis and Asterabad. It is possible that several
races of house mice (Mus musculus subsp.), which have been
introduced into the Caucasus from the north, should be included in this
group.
487
486
The Caucasian district is a climatic and landscape refuge for all the
mesophilous species, but its particular importance lies in the shelter it
provides for large game animals which have been displaced by man from
woodless piedmont plains, lowlands and high plateaus. It is a large natural
reserve in which many of the large animals have survived, and which may
in the future serve a variety of practical and scientific purposes.
The protective qualities of the mountain relief and forest vegetation
account for the great variety and high rate of survival of carnivore and
ungulate species and individuals since the Pleistocene in this region, in
contrast with the adjacent woodless plain areas.
If measures were taken to control human intervention and to develop
Suitable landscapes, they would undoubtedly promote the expansion of the
Recent ranges of mountain [Caucasian] wildcat, panther, mountain boar,
deer, roe deer and bison into the surrounding plains, thereby making the
Caucasus a new species-dispersion center.
On the basis of studies of the mammalian fauna, it is difficult to justify
any segregation of a highland, or alpine, complex and subdistrict. There
is not a single mammal connected in one way or another with the subalpine
and mountain-forest zones which does not also occur in the highland zone
of the Caucasus. The so-called alpine species — Promethean vole and
Caucasian birch mouse — occur both in alpine meadows and in the subalpine
and meadow areas of the middle mountain-forest zone. Caucasian goat,
chamois and Caucasian snow vole attain their maximum population in this
forest zone. And so one can only speak of an alpine ecological assemblage.
The true alpine species do not occur as far downslope as the lower mountain
forest belt or in the piedmont lowlands. This distribution pattern creates
the transitional phytolandscape and geomorphological characteristics
mentioned above.
The ecological assemblage of animals of the highland zone of the Greater
Caucasus is relatively poor in species: 3 insectivore, 8 carnivore (of which
5 inhabit the region only in summer), 1 lagomorph, 8 rodent and 5 ungulate,
of which 2 migrate to this zone only in summer.
In view of the Miocene age of the Greater Caucasus relief and the Upper
Miocene age of the Caucasian mountain vegetation zones, the isolation of
these forms must have begun in the Lower Pliocene. Evidently the known
morpho-physiological adaptations of highland mammals also date from the
Lower Pliocene, and reflect to some extent the orogenic and other
associated physical processes.
Caucasian highland mammals are characterized by the usual adaptations
to lower atmospheric pressure and temperature, such as higher rate of
metabolism, red blood cell count and blood hemoglobin, enlarged heart, etc.
In the jagged topography and rock debris of their environment, the fossorial
habits of goats, chamois and snow voles were weakened, and they acquired
the habits of walking and climbing on ledges and inclined rock faces. Boar,
roe deer, deer, goat and chamois developed diurnal and seasonal migratory
habits, both vertical and horizontal, in the face of such environmental
factors as bloodsucking diptera, the distribution of mineral springs and
seasonal food deficiencies which followed snowfalls. The snow vole's habit
of stocking hay undoubtedly developed because of the plentiful shelters
available in rocks where food remains dry in spite of the heavy atmospheric
precipitation.
488
487
The occurrences of some endemic highland fossorial rodents (pine vole
and Promethean vole), which feed on rhizomes, came about because of the
disappearance of tree and bush vegetation and the development of areas of
humid meadows, meadow-steppes and bare soil in the orogenic process.
The relatively poor specific composition of the highland fauna of the
Greater Caucasus, which is easily seen upon comparison with the fauna
of the steppe plateau of the neighboring Lesser Caucasus, can be accounted
for, in our opinion, by four factors:
1. The accelerated orogenesis which probably caused the extinction of
many specialized species of the Miocene mountain-forest faunal complex.
2. The lack of other nearby mountain centers of speciation, i.e., the
relatively early isolation of the area and the difficulties of new immigration.
3. The space limitations of the highland life zone imposed by the sharp
relief of the terrain: steep slopes and no plateaus suitable for carrying
animal migrations. (These features probably account for the contemporary
absence of mountain mouflon and marmot. )
4. The inability of the taiga-arctic species to colonize the Caucasian
highlands, so that the area did not benefit from migrations caused by the
European glaciation.
The vertical boundaries in the Quaternary ranges of Caucasian mammals
underwent relatively minor shifts, which were caused by the advances and
retreats of glaciers with concomitant shifts in the phytolandscape zones.
The segregation of the ranges of the endemic species mentioned above into
two sections — Caucasian and Lesser Caucasian — occurred primarily in
post-glacial time, as xerophytic vegetation developed on the Surami Range.
The Caucasian type of mountain-forest mammalian complex reached its
maximum development on the ridges and slopes surrounding the Colchis
(Rion) lowlands. This saturation of the western Caucasus with Caucasian
endemic forms cannot be linked to the Pliocene and Quaternary Pontian
landmass in the Recent Black Sea basin, as some investigators have done.
To the east, over the Greater and Lesser Caucasus, the ranges of these
ancient autochthonous elements and later Pleistocene forms, connected
with highland meadows and mountain, broadleaf forests, gradually decline;
their distribution becomes discontinuous and even shifts into other zones
and zoogeographic districts.
At present, the species composition of the mammalian fauna of the
Colchis lowlands is impoverished, particularly in rodents and ungulates.
The periodically flooded habitats are unsuitable for burrowing rodents and
paleontological and historical evidence shows that many hoofed mammals
have been exterminated by man in the present epoch.
The Caucasian district can therefore be subdivided into the western and
eastern sectors of the Greater Caucasus subdistrict (approximately on the
longitude of Tiflis and Kazbek) and the Lesser Caucasus subdistrict, based
upon the degree of saturation of the Caucasian mammalian complex. The
Lesser Caucasus is somewhat poorer in species than the Greater Caucasus,
e.g., at present, there are no Caucasian goats in this area, the distribution
range of the pine marten is narrower, etc.
Inner Dagestan and the mountain-forest area of Talysh are characterized
by the greatest faunal peculiarity. In origin, the fauna undoubtedly belong
to the Caucasian district, but, because of the pronounced changes which
took place in the original Quaternary complex, they should be categorized
in separate subdistricts.
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Dagestan Subdistrict
(Subdistrict of Dagestan hamster and Bezoar goat)
This subdistrict includes the basins of the Argun, Andi Koisu, Avar
Koisu and Samur. Its northern boundary follows the Chernye Gory on a line
between Shatyr and Buinaksk which in the west is drawn near the upper
Iora and the Krestovyi pass. The southern boundary extends from the upper
Iora along the divide to Mount Baba-Dag, and the eastern boundary is a line
drawn from Buinaksk to Mount Baba-Dag. In the Quaternary this area
was subject to uplift, and was sharply dissected by river erosion. The
presence of solpugids and scorpions may indicate that the landscapes were
dry throughout the Cenozoic and that the fauna was linked with the southern
landmasses. The phytolandscape of this subdistrict is characterized by
mountain-steppe meadows and meadow-steppes; birch groves and subalpine
steppe-meadows occur on the high plateaus. The bottoms and southern
slopes of the ravines are typically grown with wormwood formations with
an admixture of xerophilous bushes — Christ's thorn and barberry. Mountain
pine and beech forests are well developed in the upper reaches of the Argun
and Andi Koisu.
The Caucasian mammalian complex of the Dagestan subdistrict, an area
of mountain relief with a poorly developed forest vegetation, subject
to drought, is impoverished in the ancient forest and highland species.
Of 22 species of bats known in the Caucasian district, only 6 are recorded
for inner Dagestan: 2 barbastels, 1 pipistrel and 3 noctules. There are no
moles, Promethean voles or roe deer; long-tailed snow vole occurs only
along the upper Avar Koisu, and deer occur only occasionally during the
summer season. Chamois is not numerous, occurring only near the
marginal ridges in forest-covered areas, althougheast Caucasian goat is
abundant. The extinction of mouflon probably took place recently.
However, the occurrences of southwest Asian mountain xerophilous
forms — Bezoar goat and Dagestan hamster, both larger local races
existing in isolated canyons — are evidence of early (Upper Pliocene,
Middle Pleistocene) connections with southern Transcaucasia. Small races
of voles are also characteristic: Caucasian snow vole and water vole,
which, like the Transcaucasian mole vole, inhabit dry, irrigated areas.
Asterabad Subdistrict
(Subdistrict of Talysh mole and fat dormouse)
This subdistrict encompasses the iow coastal areas of the southwestern
and southern Caspian, the eastern slopes of Talysh and the northern slopes
of El'brus. Onthe Isthmus, the Lenkoran lowlands and the mountain forests
of Talysh are incorporated into a further subdivision — the Talysh sector
of this subdistrict.
The northern boundary of the Talysh sector starts at southern Mugan
(at the village of Masally) and the spurs of the Talysh near Astrakhan-
Bazar. The eastern boundary follows the seacoast from the mouth of the
Vilyazh-Chai, andthe western boundary coincides with the lower boundary
of the upland steppes at an altitude of 1,700-1,800 m. The Lenkoran
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lowlands are the youngest area geologically. They are formed of marine
sediments and sediments deposited by the rivers of the Talysh, and are
marsh-covered in winter. The phytolandscapes of the lowland and of the
mountain slopes consist of dense broadleaf, lowland and mountain forest
with some thermophilous endemic Tertiary forms, e.g., silk tree, Russian
pea-shrub and thorny liana.
Geobotanists and zoogeographers generally tend to segregate the southern
Caspian coast as a district — Asterabad — because of its Recent
isolation and peculiar features. From the standpoint of the mammalian
fauna and the paleogeography, however, the Talysh-El'brus mountain massif
belongs in the Caucasian district for the following reasons.
Even in the Lower Miocene the northern spurs of El'brus, Talysh, the
Lesser Caucasus and eastern Taurus formed a continuous coast of the
Miocene basin. The landscapes of these areas were, therefore, similar,
in spite of the influence of the Poltava flora from the east and their isolation
beginning with the Upper Miocene (Sarmatian). The common landscape
features have survived into the present. As ап example, the alder swamp
forests and mountain beech forests of Asterabad are very like, and often
indistinguishable from, the corresponding formations of Colchis.
The Talysh sector is characterized by a specific climatic and soil
regime. The mountain slopes and coastal lowland become extremely dry
in summer, although the winters bring heavy precipitation. There is
practically no subalpine zone in this sector. The upper margins of the
broadleaf forests of Talysh (oak and fruit-bearing Rosales) grade into
xerophytic, tall, herbaceous vegetation and into upland steppe or thorny
astragali formations. Because of summer drought, the compacted
zheltozem soil and lack of meadow-growth in the lowlands, there are no
moles, common field mice or pine voles. Water vole occurs only in some
marshes which are relicts of marine bays and never dry up. There are no
occurrences of common and snow vole in the forest zone; their habitat is
confined to the highest parts of the upland steppe.
As awhole, the subdistrict contains only a few survivors of the Miocene-
Pliocene Mediterranean mammals, representing the southeasternmost
limit of their distribution.
The Talysh sector is characterized by an extreme impoverishment in
Caucasian mammals, there being по occurrences of Caucasian [Persian]
squirrel, Caucasian birch mouse, Promethean vole, Caucasian snow vole,
broad-toothed field mouse, Caucasian goat or chamois. It is
possible, however, that some of these species may still be discovered on
the El'brus.
In addition to relict species of the ancient Tertiary faunal nucleus and
the widespread species of group 1 of our scheme, the Holocene mammalian
complex of the subdistrict consists of representatives of groups 2, 8 and 10.
Of group 1, the dark long-tailed white-toothed shrew is abundant in the
subdistrict, and mole and lesser shrew inhabit the upper, more humid part
of the mountain forest zone.
Chiroptera are very abundant in Talysh, occurring mainly on the lowland
near buildings and hollows. The specific composition of the bats of the
southern Caspian coast, however, is inadequately known.
The distribution pattern of carnivores in Talysh presents no new
features: fox is extremely rare in the mountain forests, as elsewhere on the
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Caucasus; small southern bear is seldom found, but otter is particularly
abundant along quiet rivers where fish are plentiful.
Rodentia are primarily represented by arboreal forms. Fat dormouse
is especially numerous from the upper margins of the forest to the sea-
coast. Black rat occurs only on the lowland, and common field mouse only
in mountain forest. Pine vole, a rare species, lives only in the upper third
of the mountain forest. Beaver is not known inthe area at present, but
might have inhabited the lowland in the Holocene. The most numerous of
the ungulates is boar. Deer were exterminated in Talysh at the beginning
of this century, but still survive in the forests of Gilan and Mazandaran.
The possibility of occurrences of bison and primitive bull has been
discussed in Chapter III.
The southern (southwest Asian) species of xerophilous landscapes (group 2)
are not numerous in the subdistrict. They include stone marten, hare and
steppe vole, immigrants from the Mugan Steppe and highland steppes, and
porcupine, which occurs from sea level to the Talysh mountain passes.
Bezoar goats do not occur in the Talysh sector, but probably lived there
in the past. The species currently inhabits the El'brus.
For historical reasons and because of specific ecological features, the
Asterabad subdistrict cannot be expected to contain any European or Eastern
European hydrophilous, forest or steppe species of Pliocene- Pleistocene
age, or any Turanian-Afghan desert forms.
Second in importance are the Holocene immigrants to the Isthmus
(group 8), of which none are definitely known among insectivores and bats,
although they probably occur, since Talysh and El'brus were affected by the
Pleistocene cooling. Reports on the Recent occurrences of jackal, striped
hyena, tiger and, possibly, lion are more reliable.
Of the accidentally introduced species (group 10), there is the Norway
rat, which at present inhabits the lowland marshes and mountain forests.
The characteristic mammalian fauna of the Asterabad subdistrict clearly
indicates that this area is typical of the mesophytic eastern Mediterranean,
and that all of its ''Indian'' elements are later additions.
The early independent development of the Caucasian mountain-forest
mammalian complex logically suggests questions on the extent of its
influence on neighboring mammalian complexes, i.e., of the southern
Russian Plain, the Transcaspian deserts and the southwest Asian highlands.
The nature and scope of this influence were determined by the high degree
of specialization of the component species of the Caucasian complex.
Their adaptations to their mountainous, humid-forest and meadow habitats
precluded most of them from migrating to the northern steppes and
northeastern deserts, and resulted in a distribution pattern which began
to develop as early as the Middle Pliocene. At that time, the extensive
landmasses formed during the regression of the Pontian basin probably
became desert under the hot, arid climatic conditions of the period of the
productive beds. Even inthe Pleistocene (the period of glaciation) the
mountain-forest species migrated only as far as the inclined piedmont plains
of Ciscaucasia, or at the farthest to the Stavropol Plateau.
The southern, southwestern and southeastern migration routes were
more feasible for the Caucasian mammals, and the influence of the
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Caucasian highland assemblage оп the alpine zone of the Armenian Highland
and eastern Turkey was quite pronounced.
Common vole, Caucasian snow vole, Caucasian birch mouse, Caucasian
deer, roe deer and even Caucasian goat could have migrated south, possibly
on a considerable scale during the major cooling epochs. Later, in the
Holocene, the effects of these migrations were blurred and in places
completely obliterated by the climate warming and the development of
xerothermal landscapes.
Of more significance was the role played by the mountain-forest area
of the Caucasian district as a transitional area on the eastward migration
routes of the European forest species. Pine marten, common red-backed
vole, common dormouse and possibly elk migrated from Europe
via the Balkans over the northern wooded slopes of the Taurus.
SOUTHWEST ASIAN DISTRICT
(District of Asia Minor hamster and gerbil)
The southwest Asian district, as understood here, implies the eastern
Mediterranean province of other authors, but excludes the Greater Caucasus
and the Crimea. It encompasses Asia Minor, northwestern Iran, the
Armenian Highland and the lowlands of eastern Transcaucasia.
The animals characteristic of this district belong to two subcomplexes —
the upland-desert and upland-steppe. These ranges replace each other
in some places, sometimes in a mosaic pattern, sometimes in altitudinal
-zones. The upland-steppe variant has no clear boundaries on the Isthmus
because its representative species (''medium" hamster [? Asia Minor
(golden) hamster], William's jerboa, Bezoar goat and other more
specialized species) migrated far to the north in the course of their
evolution, following suitable landscapes (Figure 189).
The upland-desert variant consists of narrowly specialized desert
species, which accounts for the more conservative nature of this subcomplex
and its clearly developed northern boundary. The range of the gerbils
(Persian, Asia Minor and Vinogradov's) presents the best developed
northern boundary of the upland-desert subcomplex distribution. The
boundary is more or less clearly delineated, cutting at some places into
the Sevan subdistrict along ravines and river valleys, following mountain
slopes and zones of xerophytic vegetation (Figure 190).
Most of the district is at least of Oligocene age. Remains of Oligocene
plants are known from a locality in the middle Araks valley, a deposit of
Oligocene mammals from a locality near Alkhaltsykh, Upper Miocene
mammals from the vicinity of Lake Urmia, and Pliocene mammals from
the Zanga ravine. The periphery of the district (e.g., the Kura lowlands)
is considerably younger — Upper Pliocene and Quaternary.
The predominant Recent types of landscape of the northwestern Iranian
uplands, eastern Turkey and the middle Araks valley are sun-burned
ridges of volcanic and sedimentary rocks, alternating with valleys and
narrow ravines. The mountain slopes are covered with astragali formations
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with small xerothermal shrubs, buckthorn, almond and ephedra. Sparse
juniper forests with groupings of silver hawthorn and williowleaf pear have
survived in some places. The hot broad valleys are characterized by tree-
like mountain saltwort and wormwort, and the river banks are covered by
tugai thickets of poplar (Populus diversifoli a), oleaster, buckthorn
and tamarisk.
Upland-steppe formations with sheep's fescue predominating and some
loose-bunch grasses cover the altitudinal ranges of the plateaus from
1,600-1,900 m. Alpine-type meadows occur at altitudes no lower than
2,000 m, e.g., in the region of Khvoy and Bayazet. Smalltracts of mountain
pine and occasional oak forests survived in eastern Turkey in the region ^
of Van, Kars and Kagizman, and some oak forests were preserved on the
Armenian Highland.
The mammalian complex of the district consists of species from genetic
groups 1, 2, 4, 5, and 8.
It should be noted that most of the species of group 7[? group 8], which,
further north on the Isthmus, are postglacial immigrants, in this district
are either endemics or immigrants from the south in much earlier times.
In other words, part of group 8 can be eliminated for this district, and
included in group 2.
In that part of the district that is within the U.S.S.R., the typical
mammalian complex appears in only two small sectors. The district also
includes two large sectors with mammalian complexes which underwent
considerable change.
The general characteristics and mammalian geographic features of the
sectors in Soviet territory are given below and in Table 103.
West Iranian Subdistrict
ARAKS SECTOR
(Sector of Persian gerbil and mouse-like hamster)
The northern boundary of this sector extends from the Kagizman area in
the west, follows the southern slope of the Armenian Highland and Karabakh,
crossing the Araks in the east, and enters the Iranian part of Mugan in
the vicinity of the mouth of the Akera. The southern boundary is
conditionally drawn along the Araks. This sector can be seen to include the
entire left half of the middle Araks valley with its characteristic rocky
elevations: Alindzha-Dag, Ilyanlu-Dag, Darry-Dag and Negram-Dag. The
main type of vegetation is saltwort—wormwort formations and grasses.
In the Holocene tugai vegetation grew along the river banks, but later this
was partly destroyed and partly replaced by planted groves of plane tree
and walnut.
The Holocene mammalian complex of the Araks sector consists of almost
all the species of the upland-desert variant of the southwest Asian type.
Group 1 is composed entirely of widespread species; there are no
occurrences of mesophilous Caucasian species. The predominant forms are
noctules, pipistrels, carnivores and mouse-like rodents.
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Most of the species of this group inhabit either the relict wooded areas
or biotopes of the cultivated landscape, i.e., gardens and settlements.
The faunal background is provided by 34 representative species of
groups 2 and 8 (considered as one group): long-eared hedgehog and little
white-toothed shrew among the Insectivora; horseshoe bats among the
Chiroptera; hyena, stone marten, tiger polecat and other forms among the
Carnivora; hamster and gerbil among the Rodentia; kulan, Bezoar goat,
mouflon and goitered gazelle among the Ungulata. The index species are
Persian gerbil, Transcaucasian mole vole and mouse-like hamster.
The number of species which recently became extinct is considerable:
kulan, horse, deer, goitered gazelle and dromedary (?). Their extinction
was caused by the early occupation of the area by cattle-raising tribes and
the terrain advantages of an intermontane valley for easy extermination
of large animals. The central Asian or Turanian-Afghan elements
(group 4) are poorly represented. These species include manul, Aralykh
small five-toed jerboa and, possibly, the Arazdayan gerbil. The region
of the middle Araks is the northwestern limit of their present distribution.
The postglacial group (8) includes tiger, jungle cat and jackal. Tigers
still occurred occasionally in the last century and jungle cat still inhabits
the tugai and reedlands of the Araks valley, but jackals are exceedingly
rare there. The migration of a number of Chiroptera species from the
south during the postglacial warming of the climate seems a possibility.
These species are identified as ancient Mediterranean forms. Of the latest,
casually introduced species (group 10), Norway rat is noteworthy.
UPLAND TALYSH STEPPE SECTOT
(Sector of Persian gerbil and migratory hamster)
This sector occupies the plateau in the upper reaches of the Vilyazh-Chai,
and the Zuvanda depression in the upper reaches of the Vasharu-Chai and
Vassaru-Chai near Kel'vyaz and Kelakhan. The lower boundary of this
sector coincides with the upper boundary of the forest and xerothermal high-
grass lands on the western, northern and easiern slopes of the Talysh
Range at altitudes of 1,700-1,800 m. The landscape of the plateau is
composed of gentle slopes and level ground of chestnut soil with a thin cover
of low loose-bunch grasses (brome, mouse barley and other forms)
alternating with thorny astragali and Acantholimon. The landscape
is prominently marked by the isolated peaks of Kyz-Yurda, Mount Kelakhan
and other mountains with fields of volcanic rocks.
The Holocene niammalian complex mainly consists of species of genetic
groups 2 and 5. The common species are: among insectivores, long-tailed
white-toothed shrew and long-eared hedgehog; among carnivores, fox, wolf
and badger; among rodents, William's jerboa, steppe vole, Transcaucasian
mole vole, Persian gerbil, migratory hamster and Asia Minor hamster.
Asia Minor snow vole occurs on Mount Kelakhan, which is obviously a relict
habitat. The common vole occurs on the higher parts of the plateau in
areas planted to rye. European hare is characteristically plentiful.
There are no hoofed mammals in the sector; however, kulan, mouflon
and Bezoar goat undoubtedly inhabited the area in the recent past.
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Sevan subdistrict
(Subdistrict of Asia Minor snow vole and Asia Minor hamster)
The Sevan subdistrict includes the steppe uplands of the Lesser Caucasus.
The northern boundary of the subdistrict follows the mountain passes
of the Adzhar-Imeretian ridge and curves around the northern part of the
Dzhavakhetia highland, including the Mokrye Gory, Somkhetskii and
Bezobdal'skii ridges. Farther east the boundary follows the divide of the
Pambak ridge, passingnorthof Lake Sevan, and part of the steppe uplands
of Karabakh. The southern boundary lies along the southern slope of the
Karabakh, Zangezur and Daralagez ridges at altitudes of 1,500-1,700 m.
To the west it passes beyond Yerevan on the southern slope of Alagez,
entering Turkey in the vicinity of Kagyzman.
The Sevan subdistrict landscape is high-plateau with ridges of gentle
relief and isolated peaks; the lower areas are covered by steppe with
sheep's fescue and loose-bunch grasses predominating. Canyons, fields
covered with broken rock and sunlit areas are typically grown with
astragali and dry, sparse forests of juniper, willowleaf pear and silver
hawthorn, which have been partially destroyed. Rare oak forests have been
preserved on the northern slopes of the canyons. The higher sections,
upwards of 2,000 m, are characterized by steppe meadows, and true
mesophytic meadows are developed at still higher altitudes.
The Sevan subdistrict was an area of contest between the central
southwest Asian (xerophytic mountain desert) landscape and the Caucasian
(mesophytic forest) landscape, a contest which was reflected in the Holocene
mammalian complex. The complex has no original features, but consists
of a Caucasian-southwest Asian species combination.
The boundaries of the subdistrict are best delineated by the distribution
of relatively xerophilous species of rodents, which inhabit upland-steppe
and steppe-meadow landscapes. At high altitudes their ranges extend into
Iran and eastern Turkey.
The mammalian complex of the subdistrict comprises representatives
of groups 1, 2, 5, 8 and 10; the total number of extant forms is 43.
The most completely represented groups are 1 and 2; there are no
occurrences of species of the early European forest and Eastern European
steppe fauna (groups 3 and 6). Nor are there any Turanian forms in the
complex (groups 4 and 9).
The ancient Caucasian and widely distributed species (group 1) are
poorly represented as compared with the composition of the same group
in the Caucasian district. In the Sevan subdistrict they consist of
Transcaucasian hedgehog, Caucasian mole, Caucasian shrew, pocketed
bats, noctules, southern bear, Caucasian birch mouse, Caucasian snow
vole and roe deer.
Up to the present time in the Holocene, 5 species of hoofed mammals
have become extinct: boar, deer, chamois, bison and primitive bull.
Some of these always inhabited the subdistrict, some migrated from the
forest zone of the northern slopes of the Lesser Caucasus.
With the exception of bats and carnivores, the species listed were,
and still are, inhabitants of the most mesophytic, even hydrophytic,’
biotopes on elevated parts of the subdistrict: the banks of streams, springs,
marshes and relict broadleaf forests. Lynx and bear inhabit relict oak
forests.
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Of the ancient Pliocene species of [гап1ап-Аз1а Minor or eastern
Mediterranean origin (group 2), it is those which are associated with
upland-steppe biotopes which inhabit the Sevan subdistrict. There is a
poor representation of upland-desert species: long-eared hedgehog,
Persian white-toothed shrew, horseshoe bats, stone marten, European
hare and pika (now extinct), William's jerboa, Asia Minor suslik,
lesser mole rat, Asia Minor hamster, Asia Minor snow vole,
Shidlovskii's steppe vole, Bezoar goat and mouflon. The horse and
possibly the dromedary became extinct in the Holocene.
The Asia Minor suslik and lesser mole rat now inhabit only the western
part of the subdistrict although their distribution on the plateau has
fluctuated considerably throughout the Holocene.
In some parts of the subdistrict, the ranges of species of groups 1 and
2 form a mosaic pattern which is caused by the mountainous relief.
There are no occurrences of early European forest, steppe and
hydrophilous forms and central Asian desert and semidesert species. ивы
subdistrict is inhabited by some species of northern (European?) origin
which migrated there only in the Pleistocene (group 5). These species
include the common vole (?), which inhabits the subalpine meadows, and
the Caucasian wildcat, which inhabits the relict forests. At the present
stage, it is difficult to identify any species which can be considered
postglacial immigrants (group 8), although there were probably some
belonging to various orders. It seems particularly probable that some bats
resettled in the Armenian Highland following the Upper Pleistocene cooling,
and that some southern carnivores (striped hyena, lion, tiger) began to
migrate there seasonally.
The desert ungulates — kulan and goitered gazelle — probably also
migrated from time to time to the gently sloped steppe sections of the
plateau from the middle Araks valley.
Of the latest synanthropic immigrants, it is worth noting the Norway rat
and house mouse.
The development of the ecological assemblages and mammalian complex
of the Sevan subdistrict in the Quaternary proceeded under intensive
orogenic conditions, accompanied by faulting and extensive lava flows which
covered the western part of the country in various directions.
The Pleistocene glaciation of the highlands was not extensive and did not
figure significantly in the development of the local mammalian complex.
Man's settlement, which began in the Acheulean and developed through the
Bronze and Iron Ages, was the chief reason for the disappearance of large
mammals from the subdistrict.
Within the Sevan subdistrict an isolated sector can be distinguished in
the cold Akhalkalaki highland — Satunin's ‘district of the sources of
the Kura River.'' This sector differs from the rest of the subdistrict in the
presence of lesser mole rat and of Caucasian snow vole and Caucasian
birch mouse in the Mokrye Gory mountains, which form the northeastern
boundary of the sector. The cold climate and the availability of spring
water on the Dzhavakhetian Plateau created conditions favorable for the
development of such species as water shrew and common vole. Conversely
these reasons are also accountable for the absence of steppe vole. The
naturally treeless landscape and low relief of the plateau hastened the early
displacement of large carnivores and ungulates by man.
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Kura subdistrict
(Subdistrict of steppe vole and red-tailed Libyan gerbil)
The Kura subdistrict is located between the eastern parts of the
Caucasian district and most of it coincides with the areas of wormwood—
saltwort semidesert and beard-grass steppes in the lowlands of eastern
Transcaucasia. It is shaped like a wedge extending to the foot of the Surami
ridge and bounded by the Caspian in the east.
Between Karabakh and Talysh it grades into the western Iranian
subdistrict of the southwest Asian district.
The various parts of the subdistrict differ in geologic age. The eastern
part at 20 m below sea level was probably covered by the Holocene sea;
areas lying 30-40 m above sea level were covered by the Khvalynsk, Khazar
and Baku seas. The foothills and piedmont plains of the Lesser and Greater
Caucasus and the entire valley of the middle Kura, from Mingechaur to
Gori, are of Late Tertiary age.
The landscape and geobotanical zones can be described as follows:
level areas of saltwort—wormwood semidesert; ephemeretum and beard-
grass steppes; tugai forests on the Kura, Araks and their tributaries;
reed vegetation around lakes and in floodplains; and pistachio-juniper
forests on the eroded clayey ridges (''bozdags'') of the northern slopes.
The age of this landscape is at least Lower Pleistocene, and its early stages
of formation were probably in the Upper Pliocene.
The Kura subdistrict cannot be rated as impoverished and undersaturated
in species, as zoogeographers have sometimes done because of the young
age of the Kura-Araks lowlands and their recent emergence from the sea
waters. Only the group of mammals of the southwest Asian uplands, which was
the ancient core of the developing local complex, can be considered poor
in species. And only the biotope of the open plain semidesert, particularly
the Shirvan steppe, can be regarded as undersaturated in species. The
mammalian complex consists of representatives of genetic groups 1, 2, 4,
8 and 10.
There are at least 25 species of the Caucasian mesophilous and widely
distributed species (group 1) in the subdistrict. The most characteristic
are: Caucasian mole, black rat, boar and deer which occur in relict
mesophytic habitats (tugai, oases, reed marshes, etc. ). The carnivores
are represented by widely distributed forms: wolf, little fox, weasel,
badger, otter and bear.
The nucleus of this subdistrict comprises up to 22 species of Pliocene
age (group 2).
The ranges of most of these species extend south, southeast and
southwest thereby corroborating our delineation of this subdistrict. The
index insectivore species (long-eared hedgehog, long-tailed white -toothed
shrew and lesser shrew) are characteristic of the dry biotopes of the open
semidesert.
The index Chiroptera (up to 9 species) inhabit caves and buildings and
feed in tugai areas and the margins of oases. The most commonly occurring
species in the Kura subdistrict are horseshoe bats, Mediterranean (Kiihl's)
vespertilio, tiger polecat and African wildcat, whose ranges penetrate the
Caucasian Isthmus from the south in this part of eastern Transcaucasia.
Cheetah recently became extinct.
498
497
The characteristic ranges of the southwest Asian rodents (up to 6 species)
are either relicts (as in the case of Asia Minor [William's] jerboa and
Transcaucasian hamster) or poorly developed (as in the case of Asia Minor
gerbil). The most numerous of these species are migratory hamster,
red-tailed Libyan gerbil and steppe vole.
There is a possibility that the subdistrict was still inhabited in the
Holocene by horses of the perissodactyls, and by argali of the artiodactyls.
There are no representatives of group 3 (European) in the subdistrict.
Species of Pliocene age — eastern, Turanian and Turanian- Afghan (groups
4 and 9) — are not numerous. In this respect the mammalian complex in this
area evidently differs significantly from the insect complex, particularly
the beetle complex (see Semenov-Tyan-Shanskii, 1936). Neither ancient
nor younger Turanian species of insectivores, lagomorphs, perissodactyls
and artiodactyls occur in the subdistrict.
‚Тре small five-toed jerboa, which mainly inhabits the ephemeretum
semidesert of the Apsheron and Kabristan, maybe the only representative
of the early immigrants originating in Turan.
It will be recalled that some of the species of the Russian Plain (corsac
fox, great jerboa, saiga) migrated to this area in the Middle Pleistocene.
There are only 6-7 of the southern thermophilous species which migrated
to the Isthmus in postglacial time (group 8). It is possible that this group
should include some small insectivores and bats which we have categorized
as Pliocene immigrants.
The late appearance of jackal, stripped hyena, goitered gazelle, tiger,
lion and kulan is beyond doubt; of these, the last three are completely
extinct. The accidentally introduced species which have inhabited the area
during the historical epoch include Norway rat, a contemporary inhabitant
of lake and marsh environs and of all large population centers.
Present-day agricultural activity promotes further saturation of this
subdistrict with small Caucasian species.
SUBREGION OF ВЕАБТЕВМ HUROPE AND
NORTH KAZAKHSTAN
EASTERN EUROPEAN DISTRICT
Ciscaucasian subdistrict
(Subdistrict of common hamster and striped field mouse)
The ranges of the index species of this subdistrict developed from the
north, and we have, therefore, drawn a purely conditional northern
boundary along the Manych. The western boundary follows the seacoast
from the lower Don to the mouth of the Kodor. In western Transcaucasia,
only the narrow belt of the coastal terraces over which some Eastern
European species migrated from the south falls into this subdistrict.
The southern boundary coincides at all points with the northern boundary
of the Caucasian district. The eastern boundary, starting from the lower
Kalaus in the north, follows the middle Kuma to the Terek near Mozdok,
bypassing the lower reaches of the Terek on the north and terminating at
499
(498
)
Species
Greater
Caucasus
subdistrict
European (common) hedgehog — |concolor
Erinaceus europaeus L. Martin
Long-eared hedgehog — =
Hemiechinus auritus Gm.
Eastern mole — Talpa orientalis
orientalis Ogn. Ogn.
Caucasian mole — Talpa caucasica
caucasica Sat, Sat., ognevi
Strog.
Common desman — Desmana
moschata L.
Lesser shrew — Sorex
minutus L.
Common shrew — Sorex
araneus L.
Caucasian shrew —S orex
raddei Sat,
Water shrew — Меоту$
fodiens Schr.
balearicus
Ogn., schel-
kovnikovi
Sat,
Little white- toothed shrew —
Crocidura suaveolens
Pall.
White- bellied white- toothed
shrew —Crocidura leuco-
don Herm.
Long-tailed white-toothed —
shrew —Crocidura russu-
la Herm.
Gray white-toothed shrew —
Crocidura lasia Thos.
Persian white-toothed shrew —
Crocidura zarudnyi Ogn.
Pygmy white- toothed shrew —
suncus etruscus Savi
Great horseshoe bat —Rhinolo-
phus ferrum-equinum Schr,
Mehely's horseshoe bat —Rhino-
lophus mehelyi Matschi
Southern horseshoe bat —Rhino- Jnordmanni
lophus euryale Blas,
TABLE 103. Species and geographic distribution of Holocene mammals on the Caucasian Isthmus
Eastern Mediterranean
Caucasian district
Asterabad
subdistrict
Lesser Caucasus
subd istrict
Dagestan
subdistrict
ubdistric Talysh forest
sector
Tumanicus transcaucasicus Sat,
Barr, et Нат.
transcaucasica|talyschensis
Dahl. М. Мег.
ognevi Strog. —
volnuchini Орп.
caucasicus Sat,
traddei Sat,
dagestanicus | leptodactylus
Нерг. et Form. Sat.
dinnicki Ogn,
Note; Asterisk (*) indicates species which became extinct during the last centuries and decades of the present
500
(499)
Eastern European Central Asian
subregion Е :
steppe subregion | desert subregion
Eastern European
Southwest Asian district Western district
district
: nents Northern Caspian
Western Iranian subdistrict eaten
Ret. Kura sub- Ciscaucasian SER CUSEEIOG
Sevan subdistrict ue die a
district subdistrict
Upland Talysh Terek- Kuma
Araks sector
steppe sector sector
transcaucasicus Sat, rumanicus Barr. et Ham,
calligoni Sat, brachyotis Sat, auritus Gmel.
we — transcaucasica = = ma
Dahl.
= = — caucasicus Sat, =
= = — = caucasicus Sat. я
= = raddei Sat, — — es
= = leptodactylus balearicus Ogn. =
Sat.
= = — — dinnicki Ogn. |suaveolenis Pall,
persica Thos. + +
guldenstaedti Pall. = =
zarudnyi Ogn. = = = =
= = us a ae =
+ — = +. + =
= = + + + =
+ = = + — =
era, Plus sign (+) indicates nominal forms and forms not identified closer.
501
TABLE 103 (continued)
(500)
Eastern Mediterranean
Caucasian district
Species
Asterabad
Greater subdistrict
Dagestan Lesser Caucasus
Caucasus are re
ВЕН subdistrict subdistrict
subdistrict Talysh forest
sector
Small horseshoe bat —Rhino- + _ — +
lophus hipposideros
Bechst.
Blasius' horseshoe bat — Rhino- = = = =
lophus blasii Peters,
Bechstein's mouse-eared bat — + _ = —
Myotis bechsteinii Kuhl
Tricolor mouse-eared bat — + = = =
Myotis emarginatus
Geoffr.
Sharp-eared mouse-eared bat — = + + =
Myotis oxygnathus Mont,
Whiskered mouse-eared bat — + = + +
Myotis mystacinus Kuhl
Natterer's mouse-eared bat — + + = =
Myotis nattereri Kuhl
Long-winged bat — М iniopte- + = oy =
rus schreibersii Kuhl
Long-eared bat —Plecotus wardi Thos, = + =
auritus L.
Asiatic barbastel —Barbastel- == + = ==
la darjelingensis Dobs.
European barbastel —Barbastel- + + + =
la barbastellus Schr,
Leisler's noctule —Nyctalus + = = =
leisleri Kuhl
Giant noctule —Nyctalus + = + =
siculus Palumbo
Common noctule —Nyctalus + = i} =
noctula Schr,
Pipistrel — Уезрег!!110 + + + +
pipistrellus Schr.
Nathusius' vespertilio — Уезрег- + = + +
tilio nathusii Keys. et
Blas,
Kuhl's vespertilio—Vesperti- — — - -
lio kuhlii Kuhl
Savi's vespertilio—Vespertilio|caucasicus = — —
savii Bonap. Sat.
Bicolor vespertilio—Vesperti- + = + +
lio murinus L.
Bobrinskii's vespertilio —Ves- = = — =
pertilio bobrinskii Kuz.
Northern vespertilio — Уезреги! - = — — —
lio nilssonii Keys. et Blas.
502
(501)
Central Asian
desert subregion
3 Eastern European
subregion р
steppe subregion
Eastern European
м Western district
district
Southwest Asian district
Northern Caspian
subdistrict
Western Iranian district
Kura sub- Ciscaucasian
district subdistrict
subdistrict
Upland Talysh
steppe sector
Terek-Kuma
sector
Araks sector
+ — — + + —
+ — — + — —
— — — + — —
+ — + + + —
+ — + + + —
araxenus — + = aus и.
Dahl
+ — — + — —
+ — + + — —
= == + + — —
a — — — + —
+ — — + + —
ar = + + ch —
+ - - + — -
|
+ — — + - =
= = - сацса- = -
sicus Sat.
| + - = + + =
|
| - - - - : =
= as Bs + к Le
503
TABLE 103 (continued)
(502)
Eastern Mediterranean
Caucasian district
Species
Asterabad
Greater rs subdistrict
4 Dagestan Lesser Caucasus
Caucasus geet ree
ar subdistrict subdistrict
subdistrict Talysh forest
sector
Serotine vespertilio — Уезрег- + + + +
tilio serotinus Schr,
Ognev's уезре Шо — Уезрег- = = = =
tilio ognevi Bobr.
Pocketed bat —Tadarida — — fe =
taeniotis Raf,
Jackal—Canis aureus L. + — + +
Wolf—Canis lupus L., cubanensis + +
Орп.
Кох — Уц1 рез vulpes Ц. caucasica Jinn. alticola Орп. +
Corsac fox — Vulpes согзас L. - - = =
Striped hyena — Нуаепа — — = =
hyaena L
European brown bear —Ursus caucasicus Smirn., meridionalis Midd.
arctos L. arctos L.
Pine marten —Martes lorenzi Ogn. = lorenzi Ogn. =
martes L,
Stone marten —Martes foina nehringi Sat,
Erxl,
Tiger polecat—Vormela = - - —
peregusna СШЧ.
Polecat — Putorius — — — —
putorius L.
Siberian polecat —Putorius = = = =
eversmanni Less,
Mink —Lutreola lutreola L. |turovi Kuzn, = = =
et Nov.
Weasel —Mustela nivalis Г. caucasica Barr, et Ham,
Ermine —Mustela +? = - —
erminea Ц.
Badger —Ме1ез meles L caucasicus Ogn, canescens Blanf,
Otter —Lutra lutra L. + + + +
“ Tiger—Panthera tigris L. = = = se ptentriona-
lis баг.
* Lion—Panthera leo L. — — — =
Panther —Panthera pardus L. ciscaucasicus Sat, tullianus Уаепс.
African wildcat —Felis lybica - = = =
Forst, |
European wildcat —Felis : -
: . caucasicus Sat.
silvestris Schr,
Manul —Otocolobus — — = =
manul Pall,
504
(503)
Central Asian
desert subregion
Eastern European
steppe subregion
subregion
Eastern European
district
Southwest Asian district Western district
Northern Caspian
Western Iranian subdistrict pian
subdistrict
Ciscaucasian
subdistrict
Sevan subdistrict Kura subdistrict
Terek- Kuma
Araks sector
sector
+ — — + + +
+ — — + - —
— — — — + —
+ — — + + +
+ + hajastanicus +
Dahl.
alpherakyij + kurdistanica Jalpherakyi Sat.|stepensis Braun, karagan Erxl,
Sat, Sat. caucasica
Dinn,
= = = = kalmykorum Ogn.
cs — — satunini Matschi * —
— — meridionalis Midd. SF EVO EOS Ih, —
nehringi Sat, nehringi Sat. =
+ + + + + +
— — — — +? =>
— — — — + +
fei & = _ turovi Kuzn, et =
Nov.
caucasica Barr, et Ham. dinniki Sat,
= = | — = +? +?
minor Sat. caucasicus Ogn.jheptneri Ogn.
+ — + + + +
* = — * = =
tullianus Valenc. * =
+ — — + = —
Ls pS = = caucasicus Sat, =
505
TABLE 103 (continued)
(504)
Eastern Mediterranean
Caucasian district
Species
Asterabad
subdistrict
Greater Lesser
Dagestan
Caucasus subdteerice Caucasus
Е i ИИ
subdistrict subdistrict Talysh forest
sector
Jungle cat—Felis chaus Gild, + = + +
Lynx —Felis lynx L. orientalis Sat,
* Cheetah —Ас1попух juba- = — — —
tus Schr,
European hare —Lepus caucasicus Ogn, cyrensis Sat,
europaeus L,
” Armenian pika —Ochotona = - - —
sp.
Persian (Caucasian) squirrel — + = + =
Sciurus anomalus Gild.
Little зи ИК —Citellus musicus Men. — — —
руртаецз Pall,
Asia Minor suslik —Citellus — — = =
Cire l Pusey,
* European beaver —Сазтог . — — =
fiber’L.
Fat dormouse —Glis glis L. tschetschenicus Sat, + persius Erxl,
Forest dormouse —-Dyromys caucasica ognevi Hept, pictus Blanf.
nitedula Pall, Ogn, et Turov | et Form,
Southern birch mouse — $ icista — — = ==
subtilis Pall,
Northern birch mouse —Sicista |strandi Form, = — =
betulina Pall,
Caucasian birch mouse —Sicista + + + -
caucasica Vinogr,
Small five-toed jerboa — А 11ас- - - = —
тара elater Licht,
Great jerboa —А11аста ра = = = =
jaculus Pall,
William 's (mountain) jerboa — Al- - - — —
lactaga williamsi Thos,
Little earth hare -Alactagu- = = = —
lus acontion Pall,
Thick-tailed three-toed jerboa — - = = =
Scirtopoda telum Licht.
Northern three- toed jerboa — - - = =
Dipus sagitta Pall,
Russian mole rat —Spalax = = = =
microphthalmus Gild,
506
(505)
subregion
Araks sectors
Southwest Asian district
Western Iranian subdistrict
xanthoprym- =
nus Benn,
aralychen-
sis Sat,
pictus
Upland
Talysh
steppe
sector
Sevan subdistrict
orientalis Sat,
cyrensis Sat.
xanthoprymus
Benn.
Blanf.
williamsi Thos,
tichomirovi
Sat.
caucasicus
Nehr,
schmidti Sat,
507
Kura subdistrict
Eastern European
steppe subregion
Eastern European
district
Ciscaucasian
subdistrict
+
* orientalis (?)
Sat,
Central Asian
desert subregion
Western district
Northern Caspian
subdistrict
Terek-Kuma
sector
caucasicus Ogn.
kalabuchovi
Ogn., satunini
Svirid., boeh-
mi Кгаз$,
+
daghestanicus
Орп. et Turov,
nordmanni
Keys. et Blas.
jaculus Pall.,
fuscus Орп.
dinniki Sat,
turovi
nogai
planicola Sat,
kizljaricus Sat,
fuscus Орп,
Hept.
Sat,
TABLE 103 (continued)
(506)
Eastern Mediterranean
Caucasian district
Species
Asterabad
Greater subdistrict
Dagestan Lesser Caucasus
Caucasus i. ae, oa
deer subdistrict subd istrict =
subdistrict Talysh forest
sector
Giant Russian mole rat —$ paiax — - — =
giganteus Мег.
Lesser mole rat —Spalax _ — — =
leucodon Nordm,
Black rat —Rattus rattus L. + = + +
Norway rat—Rattus norve- colchicus + -- +
gicus Berck, Lvov
Ноцзе mouse —Mus musculus formosovi Heptn., abbotti |abbotti Waterh, |tataricus
L. Waterh, Sat,
Harvest mouse — Micromys = — — =
minutus Pall,
Striped field mouse — А pode- caucasicus — = =
mus agrarius Pall. Duk,
Caucasian yellow-spotted mouse |ponticus saxatilis argiropuli Vin. =
—Apodemus fulvipectus Svirid. Кгаз5. et Arg.
Ogn,
Yellow-necked field mouse — = — = =
Apodemus flavicollis
Melch, .
Common field mouse — A pode - ciscaucasicus Орп. arianus Blanf,
mus sylvaticus L,
Broad-toothed field mouse (Asia |euxinus — euxinus -
Minor mouse) —A podemus G. Allen G. Allen
mystacinus Danf, et Alst.
Asia Minor(golden )hamster — koenigi Sat. }raddei Nehr., |brandti Мейг. =
Mesocricetus auratus avaricus
Waterh, Орп. et Heptn.
Common hamster —Cricetus + = = =
cricetus L,
Migratory (gray) hamster - Cri- pulcher Орп. + =
cetulus migratorius Pall.
Моицзе- Ике hamster —-Calomys- - — = =
cus bailwardi Thos,
Great gerbil — Rhombomys = = = =
opimus Licht,
Midday gerbil -Meriones =: — = =
meridianus Pall,
Red-tailed Libyan gerbil — = = = =
Meriones erythrourus
Gray
Persian gerbil -Meriones = a = =
persicus Blanf,
1704 508
(507)
у Eastern European Central Asian
subregion - :
steppe subregion desert subregion
Eastern European
Southwest Asian district ОИ
district
Western district
: Soh bast Norther Caspian
Western Iranian subdistrict
. Е subd istrict
жи ES Ciscaucasian
Sevan subdistrict | Kura subdistrict BL ik
subdistrict
Upland Talysh Terek- Kuma
Araks sector
steppe sector sector
we = = = т Ех
= = armeniaca = — —
Mehely, neh-
ringi Sat.
— — — + + +
+ + + + + +
tataricus Sat. abbotti tataricus Sat, |hortulanus wagneri Eversm.,
Waterh, Могат. nogaiorum
Heptn.
— — — — + +
— — — — caucasicus Duk, =
= == = = planicola =
Svirid.
= = = = samariensis Ogn.
arianus Blanf, ciscaucasicus —
Ogn.
— brandti Мег. nigriculus —
Меш.
= == = = stavropolicus =
Sat.
+ + + + pulcher Ogn. haeus Pall,
ыы ны = = res Сь
— pal. = = Bo *
= = = = = ogaiorum
Heptn.
+ — = cCaucasicus — =
Brandt,
rossicus Heptn. = — = ws
509
(508) taste 103 (continued)
Species
Tamarisk gerbil —Meriones
tamariscinus Pall,
Asia Minor gerbil—Meriones
blackleri Thos,
Vinogradov's gerbil —Meriones
vinogradovi Heptn.
Arazdayan gerbil —Meriones sp.
Steppe lemming — La gurus
lagurus Pall,
Northern mole vole —Ellobius
talpinus Pall.
Transcaucasian mole vole —
Ellobius lutescens Thos,
Promethean vole(long- clawed
mole vole — Prometheomys
schaposchnikovi Sat,
Water vole —Arvicola
terrestris 1.
Asia Minor snow vole — Micro-
tus nivalis Mart,
Caucasian snow vole —Microtus
gud Sat.
Long-tailed snow vole —
Microtus roberti Thos,
Pine vole —Microtus majori
Thos.
Common vole —Microtus
arvalis Pall,
Steppe vole — Microtus
socialis Pall.
Common red-backed vole —
Clethrionomys glareolus
Schr,
Porcupine — Hystrix leucura
Sykes
* Kulan (Asiatic wild ass) —
Equus hemionus Pall.
* Tarpan—Equus caballus
gmelini Ant,
>
Стеатег
Caucasus
subd istrict
ognevi Tur,
rufescens
Зат., turovi
Ogn.
loginovi Ogn.
nenjukovi
Form., gud
Sat.
personatus
Ogn.
colchicus
Schidl., cis-
caucasicus
Орп.
Eastern Mediterranean
Caucasian district
Lesser Caucasus
subdistrict
Dagestan
subd istrict
— +
djukovi Ogn. persicus
et Form.,
kuruschi
Heptn, et Form.
trialetcus
Schidl,
lghesicus +
Schidl., ose-
ticus Schidl.
+ +
intermedius
Schidl.
daghestani-
cus SchidL
macrocranius Ogn., gudauri-|transcaucasi-
cus Ogn.
* ponticus
Thos,
cus Ogn.
510
Asterabad
subdistrict
Talysh forest
sector
de Fil,
schelkovni-
kovi Sat,
hirsutirostris Brandt
(509)
Central Asian
desert subregion
Eastern European
subregion ;
steppe subregion
Eastern European
district
Southwest Asian district Western district
: р Northern Caspian
Western Iranian subdistrict
: 5 subdistrict
8 te sy thls Ciscaucasian
Sevan subdistrict | Kura subdistrict ТЕ:
subdistrict
Araks Upland Talysh Terek-Kuma
sector steppe sector sector
ciscaucasicus
7h 7 8 У iD Sat.
bogdanovi Heptn. = bogdanovi = +
Heptn.
+ — — — — —
+ ae he a une =
— — — — + —
— — = = + +
+ + = — - —
persicus de Fil. caucasicus +
Ogn., cuba-
nensis Ogn.
— = satunini = = =
Schidl.
= mystacinus|transcaucasi- = macrocranius
de Fil. cus Ogn. Ogn.
+ + schidlovskii |binominatus parvus Sat,
Arg. ЕПегт.
* * * % * ыы
— —= = = * *
511
1704
TABLE 103 (continued)
(510)
|
}
Eastern Mediterranean
Caucasian district
Species
Asterabad
Dagestan Lesser Caucasus subdistrict
subdistrict subdistrict
Greater
Caucasus
А
subdistric Talysh forest
sector
= Southwest Asian horse —Equus
caballus L (subsp.)
Boar —Sus scrofa L. attila Thos.
Red deer —Cervus ela- maral Ogilby
phus L.
* Elk—Alces alces L. * caucasicus
N. Ver.
Roe deer —Capreolus capreolus L., pygargus Lyd.
capreolus L.
Saiga — ба1ра tatarica L.
Goitered gazelle — Gazella-
subgutturosa Guld.
Chamois —Вир1сарга rupi- caucasica
capra L.
West Caucasian goat—Capra caucasica
caucasica Guld. Guld., severt-
доу! Menzb.,
dinniki Sat.
East Caucasian goat — Сарга
cylindricornis Blyth
Bezoar goat (wild goat or ibex) —
Capra aegagrus Erxl.
Armenian mouflon —Ovis
gmelini Blyth
* Caucasian bison —Bison caucasicus Sat,
bonasus L.
* Primitive bull —Bos
primigenius Boj.
the Caspian coast near Makhachkala (Figure 195).
The various sections of the subdistrict differ in geologic age. The
youngest are the Kuban and Manych plains and the lower reaches of the
Terek and Sulak; the oldest are the Stavropol Plateau and the sloping
piedmont plains, which are made up of Maikop (Oligocene), Upper
Sarmatian (Miocene), Apsheron (Pliocene) and Quaternary formations.
The primeval postglacial landscapes are diversified in their features and
in their origins. To the north in the Manych area, sheep's fescue and
wormwood steppes predominate; the piedmont plains in the western area
are characterized by steppe meadows, mesophytic meadows and forest
steppe with islands of broadleaf forest and tugai-type forest growing along
512
(oh)
Eastern European Central Asian
steppe subregion desert subregion
subregion
Eastern European
т Western district
district
Southwestn Asian district
Northern Caspian
Western Iranian subdistrict И
subdistrict
Ciscaucasian
Sevan subdistrict | Kura subdistrict
bdistrict
Araks Upland Talysh а Terek-Kuma
sector | steppe sector sector
ы * * ? = —
attila Thos. attila Thos.
— — + maral Ogilby
= = * = * caucasicus =
N. Ver.
— — capreolus L. = * pygargus Lyd. =
* * р + % %
+ Е + == = =
armeniana Nas. — = Be
the river floodplains. Steppes developed on a considerable scale on the
Trans-Kuban and Terek-Sunzha plains during the xerothermal phase of the
Holocene. At the beginning of the present epoch, forests began to cover
the steppe areas (Zakharov, 1935). The typical landscape of the
southeastern area is dry wormwood steppe which grades into saltwort
semidesert. Thickets of reed, narrowleaf cattail and bulrush grow on the
shores of the estuaries along the Azov and Caspian coasts and on the
floodplains of the lower Kuban, Chelbas, Yeya, Terek and Sulak. These
Swampy areas are separated by sand spits covered with dry steppe
vegetation and inhabited by jerboas, hamsters and hares. Tugai forests,
alternating with plumegrass and licorice, remained intact only along the
lower Terek and Sulak.
213
1704
510
511
The mammalian complex of the subdistrict is not homogeneous:
it comprises elements of genetic groups 1, 2, 3, 5, 6 and 8, of which groups
5 and 6 furnish the foundation of the complex.
The Caucasian Pliocene group is poor in species, consisting of European
(common) hedgehog, Caucasian mole, lesser shrew, noctules and
vespertilios, widely distributed carnivores, small rodents and ungulates
(of which the only survivor is the boar). Caucasian bear, European beaver,
Caucasian deer [maral: subspecies of red deer] and Caucasian bison became
extinct in historical time.
There are 15 southern species, i.e., group 2, originating in southwest
Asia: long-eared hedgehog, white-bellied white-toothed shrew, horseshoe
bat; stone marten and tiger polecat among the carnivores; and a number
of xerophilous rodents — medium hamster, steppe vole and others. Most
of the species of this group inhabit the xerophytic biotopes of the foothills
zone.
The ancient Eastern European species (group 3) are rare; they include
desman and mole rat.
All the other extant species are later — Pleistocene and Holocene —
immigrants to Ciscaucasia, primarily from the north: from the forests
of Western Europe and the steppes of Eastern Europe.
Group 5 consists of 9-11 Pleistocene-age species of northern origin and
characteristically of forest and mesophilous biotopes: Caucasian shrew,
mink, Caucasian wildcat, harvest mouse, striped field mouse, Samarian
subspecies of yellow-necked field mouse, common hamster. Of those
species which became extinct in historical time, the following are
noteworthy: European brown bear, Caucasian elk, Caucasian large roe
deer and, possibly, primitive bull.
The ranges of most of these species are close to the Caucasian foothills.
The principal proof of their Pleistocene age on the Isthmus rests on their
poorly-developed southeastern distribution, geological data and the history
of the phytolandscapes. Since all of these species are inhabitants of
relatively cold environments, it is doubtful that they could have migrated
to the hot, arid south in pre-Baku time. The distribution of the Eastern
European subspecies of yellow-necked mouse, which, so far as is known,
is corfined to the tugai sections of the Sulak, is somewhat enigmatic.
The steppe species of Eastern Europe (group 6) migrated to the
subdistrict mainly during the Pleistocene. They are: lesser shrew, corsac
fox, Siberian polecat, little suslik, southern birch mouse, great jerboa,
northern three-toed jerboa, thick-tailed three-toed jerboa, steppe lemming
and northern mole vole.
Marmot and tarpan are among the recently extinct species.
Some of these species (e.g., little suslik) invaded the upland steppes on
the northern slope of the Caucasus; others (great jerboaand saiga) migrated
in the Middle Pleistocene by way of the Caspian coast as far as eastern
Transcaucasia.
The postglacial immigrants (group 8) comprise carnivores,
perissodactyls andartiodacyls, andinclude jackal, jungle cat and such
recently extinct forms as striped hyena, kulan and goitered gazelle. It is
possible that lion, panther and cheetah inhabited the subdistrict in
historical time.
The ranges of jackal and jungle cat within the subdistrict form a narrow
strip along the river valleys and foothills of eastern Ciscaucasia.
514
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512
The accidentally introduced species of the historical epoch are
represented by Norway rat, which inhabits the floodplains of the Don,
Kuban, Sulak and Terek, and is rapidly invading every settlement.
To the east and northeast, the subdistrict adjoins the semideserts and
deserts of central Asia which are characterized by a mammalian complex
better adapted to xerothermal conditions.
SUBREGION OF CENTRAL ASIAN (DESERTS
WESTERN DISTRICT
North Caspian subdistrict
TEREK-KUMA SECTOR
(Sector of tamarisk gerbil and midday gerbil)
This sector encompasses the semideserts and deserts of the lower
Kuma-Terek-Sulak interfluve. Its western boundary starts in the north at
the Manych-Sarykamysh divide and continues southeasterly to the mouth of
the Sunzha. The southern boundary crosses the lower Terek and Sulak and
terminates on the Caspian coast near Makhachkala. The sector corresponds
to that part of the Isthmus which was covered by the sea during the last
major Khvalynsk transgression. Its northern half is made up of marine
sediments of glacial time and is a level sandy-clayey semidesert. The
southwestern third is in sandhills; the southeastern is composed ofalluvial
loams and clays of postglacial time. The youngest areas geologically are
the northern half and the southeastern third, which were partly covered by
the sea even during the minor transgressions of the Caspian in historical
time. Most of the Terek-Kuma semidesert and desert is covered by
wormwood-—saltwort vegetation; the sandhills are covered by lopsided
oat, wormwood and sweet clover; and the coasts of the Caspian estuaries
are covered with reed thickets alternating with weeds on low elevations.
The nucleus of the Holocene mammalian complex of the sector is formed
by group-6 and group-7 species; the complex is completed by species of
groups 1, 2, 4, 8 and 9.
Group 1 is represented only by Palaearctic ubiquists which inhabit the
interzonal biotopes; mesophilous Caucasian species of group 1 do not occur
at all in this sector. Common hedgehog, wolf, weasel, badger, steppe
mouse and boar inhabit reed-grown areas and hollows between sand-
hills. These species play no significant role in the biocenoses of this
sector, however,
The only eastern Mediterranean xerophilous mammals (group 2) are the
widely adapted species: long-eared hedgehog, white-bellied white-toothed
shrew, tiger polecat, European hare, migratory hamster and steppe vole.
The Russian mole rat has been replaced by giant Russian mole rat which
occurs in isolated colonies in hollows between dunes. Small five-toed
jerboa represents the ancient central Asian species (group 4).
516
513
The Eastern European steppe species (group 6) and Turanian species
(groups 7and9), which inhabited the sector during the Pleistocene
regressions of the ancient Caspian and in the Holocene, are represented
by corsac fox, little suslik, jerboas and midday and tamarisk gerbils.
Great gerbil, manul and tarpan became extinct in historical time, but
saiga has survived in this sector.
The central Asian elements in this complex are comprised essentially
of gerbils. Tarpan inhabited this sector longer than the Ciscaucasian
subdistrict.
The southern, postglacial species (group 8) probably penetrated this
sector from the south, andthose from the northeast from centralAsia. It can
be assumed that the sector was inhabited in the Holocene by almost all the
group-8 species which occurred in the Ciscaucasian subdistrict. Of these,
only jackal and jungle cat survived until recent time.
A study of the mammalian fauna of the Terek-Kuma sector shows that
it is essentially a part of the Aral-Caspian (central Asian) deserts, and
not a part of the Caucasus zone. The animal assemblages of the sandhills
in particular resemble the central Asian assemblages, e.g., the assemblage
on the phenomenal Kumtorkala barchan* in the foothills of Dagestan.
The fluctuations in the sea level of the Quaternary Caspian affected the
development of the mammalian complex of this sector more than that of
any other Caspian area. This factor, as well as the bifurcation of the lower
Volga, strongly influenced the migration of hibernating species from the
east in the Quaterrary.
This study of the faunal influences of adjacent territories and of the
characteristics of the districts, subdistricts and sectors of the Caucasian
Isthmus, based on the history of species distribution and paleogeographic
data, reveals only a small part of the biological processes which took place
during a time of complex change in the Cenozoic terrestrial environments
around the Black and Caspian seas.
The geochronological continuity of the faunal influences, which we have
noted, will certainly be better understood as paleontological data is
accumulated and processes of speciation are analyzed. Nevertheless,
the scheme presented here is justifiable at the present stage of
zoogeographic knowledge.
The characteristics of the mammalian fauna given for each zonation
also show the degree of its saturation with species. This opens avenues
leading to new deductions on the restoration and enrichment of the fauna.
The complete list of species (and subspecies) of the Holocene mammalian
fauna of the Caucasian Isthmus is given in Table 103.(p. 500).
The effects of both casual and purposeful anthropogenic influence on the
latest evolutionary patterns of mammalian ecological assemblages of the
Isthmus, and on the changes in boundaries of the faunal complexes, are
discussed in the next chapter.
* [A symmetric dune with crescentic ground plan; gentler slope facing the wind on the convex side, and
steeper slope on the concave or leeward side. ]
517
514
515
Chapter VI
ANTHROPOGENIC CHANGES IN MAMMALIAN
ECOLOGICAL ASSEMBLAGES AND RANGES
IN VARIOUS ZONES
Man's influence on the mammalian fauna of the Caucasus can be traced
from Acheulean time by means of the kitchen middens of primitive tribes
Here, as in other regions, the Paleolithic tribes were largely responsible
for the extinction of а number of mammals and the reduction in range of
others. The extent of the human influence on different species is difficult
to gauge. The game of prehistoric hunters consisted largely of the widely
distributed and most accessible species. For example, the chief game of
Paleolithic tribes on the Trans-Kuban Plainwas bison, followed by horse,
mammoth, red and giant deer and cave hyena. On the Black Sea coast and
and in central Transcaucasia, cave bear was the chief game, followed by
boar and red deer. Bison and horse were hunted in the Lesser Caucasus
highlands. Over the millennia, the species and their proportions in the
game of Paleolithic hunters probably changed considerably from season to
season. Many species of small animals, birds, reptiles and fish used for
food may not be preserved in the paleontological record; they may well
have been consumed immediately at the hunting site or at temporary camps.
Some of the animal species used for food by primitive Upper Pleistocene
tribes in. various parts of the Caucasus are given in Table 104. The
proportions given probably relate more to the type of landscape than to
the age of the site.
TABLE 104. Number of remains (in %) of predominating species of mammals at various Paleolithic sites
in the Caucasus
Lesser Caucasus
upland
(Zurtaketi )
Northwestern Black Sea
Caucasus coast
(Il'skaya ) (Akhshtyrskaya )
Colchis
oh abe
aye (Sakazhia )
In post-Paleolithic times, man's influence on the Caucasian mammals
and their assemblages assumed many aspects:
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516
1) direct extermination of game species by unrestricted hunting;
2) indirect influence on single species, their assemblages and ranges
through changes in landscape, macro- and microclimate, soil and other
ecological features caused by industrial and agricultural activity;
3) accidental introduction of synanthropic species;
4) planned changes in ecological assemblages effected by introducing
and raising new species under natural conditions, breeding and protecting
selected species and exterminating harmful species.
9723 4 Som
FIGURE 196. Bronze arrowheads and darts used in hunting and war from Late Bronze Age
(darts) and Scythian-Sarmatian culture (Caucasian Museum collection)
As human population and dispersal areas grew and culture developed in
post-Paleolithic time, the direct influence of man on the large wild
mammals of the Caucasus increased rapidly.
The emergence of domesticated animals could not offset the magnitude
of these effects. The decrease in the number of remains of wild animals
which can be observed in post-Paleolithic cultural layers from Early to
Recent is essentially the result of the development of animal husbandry,
which is not to say that wild animals were hunted on any smaller scale
(Table 105).
The direct extermination and population decrease of a number of game
animals of the Caucasus during the last millennia was closely related to
the history of the Caucasian tribes and peoples, their migrations and wars,
and the development of their material culture, particularly weaponry.
519
517
TABLE 105. Number of bones of wild and domestic animals in post-Paleolithic localities, settlements and
ritual sites of the Caucasus and the Russian Plain
*
Percentage of
animal bones
Developed cattle raising; abundance of
81.05 mountain-forest animals, killed with spears,
bows and arrows, firearms.
Abundance of domestic cattle and wild
Monument and its age Description
Dzuars of North Ossetia. Digorized
cave, 14th-19th centuries A.D.
Khazar fortress Belaya Vezha (Sarkel)
on the Don, 8th-13th centuries A.D. 74.0 steppe and forest animals, killed in safari-type
hunt and by individual hunters.
Settlement near Baku Bayon the Apsheron, 79.4 Abundance of domestic cattle, goitered
9th- 13th centuries A.D. gazelle, seals, migratory birds; developed
hunting of goitered gazelle and kulan in
game drives; seal hunting.
Developed raising of low-grade cattle;
hunting more important as the town declined
at the beginning of the present era.
Primitive cattle-raising; abundance of steppe
ungulates,
Developed cattle-raising; hunting, including
seal hunt, important.
Developed dolphin hunting; cattle-raising
and hunting of forest animals,
Abundance of large forest animals in the
vicinity of the cave.
Ancient town of Semibratnoe, Kuban
estuary, 4th century B.C. - 1st century
А. О.
Mud-hut settlements near Tsimlyanskaya
on the Don, first millennium В.С.
Settlement near Kayakent in Dagestan,
second millennium B.C.
Mud-hut settlements near Gelendzhik
Bay, third millennium B.C.
Akhshtyrskaya cave, hunting site,
Neolithic and later beds
* Unpublished author's materials.
** Percentage of bones of wild animals from Sarkel was artificially increased in the process of sorting for
final identification.
Although we have plentiful archaeological and documentary materials
on the Caucasus, it is difficult to trace the details in the evolution of hunting
from the Paleolithic to the Recent. Ancient drawings on rocks in Kabristan
and drawings on vessels from the Trialet and Maikop burials indicate that
bows were widely used in the second and first millennia B.C.
Spears and bows were the chief weapons in the big-game hunting of the
Koban, Colchis and later tribes (Figure 196). It is characteristic of
medieval weaponry in the northern Caucasus that the iron arrowheads
and darts used in combat and the hunt show remarkable variations in type
(Figure 197).
The horse-mounted big-game drive was the principal hunting method
used by the nomads of the Isthmus plains during the last three millennia,
sometimes with either permanent or temporary fences or traps placed
across gulleys and river valleys.
In addition to group hunting, individual game hunting was widely
practiced in the forests and mountains.
520
a + 9 ст
FIGURE 197. Iron arrowheads and darts from the Middle Ages — 7th- 9h centuries A.D.
(Caucasian Museum collection)
The sharp decrease in the large-animal population on the Caucasus
undoubtedly occurred during the Middle Ages when the techniques of bow-
manufacturing and forest and mountain hunting were at a very high level.
Large-scale hunting into late medieval time was made possibly by the
conditions of a feudal society and by the existence of large bands of free
armed men which provided the necessary manpower.
Rashid ad-Din, the Iranian chronicler of the 14th century (1946 edition),
records a story of the expedition to Aladagh made by the ''king of Islam, "
Ghazan-Khan (one of the Mongolianrulers ofIran). Inthe winter of 1301-02,
Ghazan-Khan camped at Belesuvar (inthe southern Kura-Araks lowland) and
Khamashakhre, and from there he hunted inthe mountain district enroute to
Talysh and Ispakhbad.
518 "Ghazan-Khan ordered the construction of two wooden fences in the
mountains, each fence the length of one day's travel, which together would
form a wedge fifty gyaz wide at the narrow end and one day's travel
apart at the wide end. At the dead end the fences were to be closed
off as acorral. After this the warriors drove the game — mountain buffalos,
dzhurs, wild goats and asses, jackals, foxes, wolves, bears and other
various wild and predatory beasts — between the fences until all were in the
corral. The kingofIslam was seated with Bulugan-Khatun on the stage
which was built in the middle, and enjoyed the sight of the animals. Some
were killed and some set free" (pp. 188-189).
From the latest Iranian chronicles, studied by Petrushevskii (1949,
рр. 292-293), it is known that in the 16th-17th centuries in Iran, peasants
were often called for a hunting duty — ''shikari."'
"Hunter-emirs assembled on the Shah's orders with the gentry
("mulyadzins'') and men-at-arms ("nukers''), often as many аз ten thousands
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519
ог more. Theyenclosedthe large area selected for the hunt to prevent the
animals from escaping. The circumference of the enclosure gradually
became smaller asthe hunters moved intothe area. The circumscribing line
was called ''dzherke,"' 'negre" or "Котагра" (р. 292).
During the Shah's hunt in Mian-Kaleh in the forests of Mazandaran in
1028 (1618 А.Ш. — М. У. ) the ring surrounding the animals was 3 farsangs
(18-20 km — М. У. ) long and 2/3 farsang (4-4.5 km — N.V.) wide. Before
the beginning of the big hunt, the animals were driven for several days
into the encircling ring. Hundreds and even thousands of large animals
were killed during such hunting. In addition to the mounted warriors, foot
soldiers were used to drive the animals. During the Shah's big hunt in
the forests and thickets of Gilan (southern Caspian coast —N.V. ) in the
spring of 1002 (1564 A.D. —N.V.) nearly 10,000 peasants from among the
people of Gilan were called to drive the animals into the dzherke" (р. 293).
During the Middle Ages, kulan and goitered gazelle were hunted in the
middle Araks valley, and forest animals in northern Armenia and Georgia
on a similarly grandiose scale, as described by Ananiah of Shirak in the
"Book of Problems" (see Ter-Pogosyan, 1947), in Rustaveli's (1937) and
Nizami's (1940) poems and in Prince Vakhusti's ''Geography" (1904).
The large population of ungulates had already been considerably
decreased by primitive hunting by the time firearms were discovered,
introduced and perfected in the Caucasus.
The perpetual wars between small tribes and feudal lords occupying
separate canyons and the local custom of blood revenge greatly
accelerated the distribution of firearms, making them also generally
available for hunting purposes during the last centuries.
With the decline in the animal population toward the 18th-19th centuries,
large-scale hunting with fences and corrals was no longer feasible. But the
introduction of small-gauge flint and percussion-type shotguns and rifles
created new possibilities for individual hunting.
The Caucasian wars of the 19th century also contributed to the decimation
of large animals.
The final stage in this development was the introduction of the four-
barreled Berdan rifle and, later, of the Mosin rifle — the Russian three-
barreled rifle. The latter model created almost unlimited possibilities
for shooting mountain ungulates from great distances. Cattle-herding
from one pasture region to another, the arming of shepherds, and the
shipment of large quantities of arms into the mountains during the Civil
War (1917-1922) brought about the final stage in the destruction of the
Caucasian populations of deer and bison.
The evolution of the distribution ranges of individual species of
Caucasian mammals which is reviewed in Chapter IV shows the contributory
role of man in the extinction of at least 9 species in the last thousand years
(Table 106).
In addition to the forms mentioned above, deer, roe deer, goitered
gazelle, saiga and other species have declined in population and distribution
range (Vereshchagin, 1947).
Human activity indirectly affected the fauna of the Caucasian Isthmus
in very complex and diversified ways through:
1) destruction of tugai, foothill and mountain forests by cutting, fires,
cattle grazing and cultivation;
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520
2) plowing and cultivation of steppes and semideserts, accompanied
by irrigation and planting of groves, gardens and shelterbelts;
3) cattle driving which resulted in large-scale thinning of forests,
mountain slope erosion and changes in the plant formations of mountain
meadows, steppes, semideserts and deserts;
4) construction activity and development of mud-hut settlements, villages
and larger towns with buildings of various types;
5) building of roads and railroads.
TABLE 106. Chronological order of disappearance (extinction) of some Caucasian mammals
Probable time of
Species :
disappearance
Area of last habitat
Lion 10th century Eastern Transcaucasia
Primitive bull 12th century Western Ciscaucasia
Kulan 13th century Eastern Transcaucasia and (until the 18th century)
eastern Ciscaucasia
Cheetah 13th century Eastern Transcaucasia
Beaver End of 19th century Colchis and Trans-Kuban Plain
Elk Beginning of 19th century | Ciscaucasia
Tarpan 1880's Eastern Ciscaucasia
Bison 1920's Western Caucasus
Tiger 1930's Talysh
The formation of the secondary, or cultivated , man-made landscape
caused the animals to develop new behavioral patterns and new assemblages
adapted to the new conditions. These changes in the fauna followed the
development of tribal economic life, cattle-raising, agricultural technology
and industrialization. The relationships between species numbers within
small-mammal assemblages (rodents and insectivores) were affected by
particularly complex factors: the population dynamics from season to
season and over long periods as they related to changes in biotopes brought
about by a variety of agricultural techniques, e.g., crop rotation, etc.
(see Qbolenskii, 1935; Polyakov, 1950, and others).
The direct and indirect effects of human activity on individual species,
their ecological assemblages and faunal complexes is discussed below by
geomorphological and phytolandscape districts with examples.
STEPPES AND SEMIDESERTS OF CISCAUCASIA
The typical primeval postglacial landscape of the Kuban Plain, lower
Don, Manych area and steppe margins of the Stavropol area was one of
motley-grasses—sheep's fescue—feathergrass and sheep's fescue—
feathergrass steppes of the northern and southern type (Shiffers, 1953).
The river valleys and steppe gulleys were developed with shrubs and
leafy forests.
From the time of the Neolithic, the floodplain terraces were steadily
settled by cattle-raising tribes. Mud-hut settlements and towns were
523
quite common along the rivers in the Bronze and Iron ages (see Gorodtsov,
1935; Pokrovskii and Anfimov, 1937; Artamonov, 1937; Goretskii, 1948).
The spurs of the Taman Peninsula were widely colonized by the Greeks
as early as the 6th century B.C. (Kallistov, 1949). The postglacial
mammalian assemblages of the Ciscaucasian steppes were rich in species.
The floodplain forests and reed thickets on the Don and Kuban were inhabited
by wolf, bear, otter, European wildcat, panther, beaver, water rat, boar,
roe deer, deer, elk, tur and bison. The steppe proper was inhabited by
tarpan, kulan, saiga and goitered gazelle. The small-mammal assemblage
of the virgin steppes of the Manych region included common
and long-eared hedgehog, fox, corsac fox, Siberian polecat, marmot,
little suslik, hamster, jerboa and mole rat.
It can be inferred from a comparison with the Recent and from historical
data for adjacent regions (Barbaro, 1836 edition; Mikhail Litvin, 1890;
Beauplan, 1832) that the disappearance and displacement of wild horse,
kulan, saiga, tur and bison from the Ciscaucasian steppes were well
advanced as early as the Middle Ages, brought about by domestic cattle
herding and game drives by thousands of mounted Khazars, Polovtsy
[Cumans] and Mongolians.
Bear, beaver, boar, deer, roe deer and elk were displaced from the lower
Don, the Kuban, the Terek andthe Sulak by the heavy deforestation by man
during the last centuries (Kondrat'ev, 1885-86; Bogachev, 1918; Flerov
and Balandin, 1931).
It is known that extensive forests existed on the right bank of the Kuban
as late as the 18th century (Mishchenko, 1928; Rogovskii, 1928; Zakharov,
1935).
Rogovskii's map indicates nine isolated forest islands on the right bank
of the Kuban where according to the 1775-76 map of the Kuban region an
almost continuous zone of forests and bush extended from the source of
the Protoka to the mouth of the Laba.
Flood control in the deltas of the Kuban, Terek and Sulak (building of
embankments and filling the gaps in the banks) have noticeably affected the
landscape since the middle of the last century. These measures only
drained the marshes temporarily, since the river levels continued to rise.
The planned drainage and reclamation for wheat and rice cultivation of
the Kuban plavni only began in the 1930's and brought about the greatest
changes in the assemblages of rodents and carnivores.
521 Mink, otter, European wildcat and water rat were displaced from vast
areas of drained swamps and lakes during reclamation, and their habitats
were occupied by polecat, hare, common vole, steppe mouse, common
hamster and jerboa.
Military considerations originally prompted the burning and cutting of
the tugai forests on the Kuma, Terek and Sulak which began in the second
half of the 18th century during the construction of the Caucasian
fortifications (Popko, 1880; Potto, 1912). The establishment of Cossack
settlements and posts brought about a rapid extermination of large animals,
since the Cossacks were mainly engaged in hunting and fishing. During the
reign of Anna Ioanovna, it was the duty of the Grebenskie Cossacks to
supply live boars, deer, ibex (''steinboks'') and bison to the menageries
of the capital.
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522
The destruction of ''bairak'' [small-gulley] forests scattered оп the
northern slopes of hills and in the gulleys of the Ciscaucasian plains was
similar to the process described for the steppes of the Russian Plain
(Kuznetsov, 1896; Tanfil'ev, 1896; Lavrenko, 1939). Cattle breeding and
agriculture on the Ciscaucasian plains played an important part in the
formation of the ranges, populations and assemblages of small mammals
and completed the process of displacement of corsac fox, kulan, tarpan
and saiga which began in the first half of the last century during the building
of the Cossack settlements. The extermination of the bairak forests brought
about a decrease in the area of the habitats of mesophilous insectivores
(shrews, moles), smallcarnivores (stone marten, badger), and forest rodents
(forest dormouse, common and striped field mouse). The distribution of
these species on the Ciscaucasian plains gradually split into small isolated
colonies.
Examples of the residual, disrupted ranges of these species can be
observed in the Petrovskoe and Aleksandrovskoe Districts of the Stavropol
Region, on the middle and upper Kalaus and on the Stavropol Plateau.
The depletion of wolves, susliks and mouse-like rodents in the
Ciscaucasian steppes dates from the early 20th century, but planned
extermination has been carried out only under the Soviet regime. The 20th
century also marks the beginning of the latest phase of development of the
cultural landscape, which is well presented in the extensive statistics of
the collection ''The Northern Caucasus after Districting'' (1925).
During recent decades the introduction of new crops and irrigation
and melioration projects (e.g., construction of canals on the left banks
of the Malka and Kura, diversion of the Kuban waters to the Yegorlyk, and
drainage of the Kuban plavni) resulted in accelerated agriculture and an
increase in cultivated areas. Planting of shelterbelts started in the 1920's,
at first along the railroads and later, in the 1930's, in the vast croplands.
The largest shelterbelts are in Krasnodar, Tikhoretskaya and Salsk, where
they are many hundreds of kilometers in length.
The highly diversified ecological features and potential adaptations of
various species to new conditions is readily seen in the nature of the
relationships which developed between the mammals and the biotopes of the
cultural landscape.
The animal world of this region was affected by cattle raising for a longer
time than by agriculture. The destruction of the steppe-grass stand by
cattle over the centuries resulted in the replacement of motley-grass—
sheep's fescue—feathergrass steppe formations by sheep's fescue—
feathergrass vegetation, and later by sheep's fescue—wormwood vegetation.
Concomitant with this process, weed groupings developed near resting and
watering places, along the routes of cattle drives and around settlements.
As early as the 1880's extensive grain farming began to replace the fine-
wooled-sheep breeding of Ciscaucasia, from the virgin mixed-grass steppes
of the Kuban and Terek and the Stavropol area to the zone of sheep's
fescue—feathergrass steppes of the Kuma basin (Rozhdaev, 1925). Cattle
grazing of virgin vegetation cover and overgrazing and manuring promoted
the rapid expansion of suslik and jerboa ranges. The extension of the suslik
range southward from the Manych area was also promoted by plowing of
the steppe, construction of bridges over canals and rivers and established
525
523
routing of cattle drives (Sviridenko, 1927; Formozov, 1929). The
overgrazed, manured areas gave susliks and jerboas the advantages of
a good field of vision, fresh green food, consisting of mixed weeds and
grasses, fast movement and protection from predators. Both abandoned
and occupied cattle stations were concentration sites for white-bellied
white-toothed and lesser shrews and steppe voles.
Recent complexes of small-mammal habitats in the cultural landscape
of the Ciscaucasian steppes — Obolenskii's permanent or temporary ''farm
habitats'' (1935) — typically consist of pastures, roads of various types,
cultivated lands and settlements.
Each of these categories comprises a number of biotopes, determined
by the agricultural and construction methods of man and also by the
ecological adaptability of the mammal species.
Pastures. When the cattle load is light, the pastures preserve
almost all the natural biotopes of the primitive ecological assemblage
consisting of little suslik, small five-toed and great jerboas, little earth
hare, common mole rat, steppe vole, steppe lemming and mole vole.
Pastures located near settlements, particularly if the soil is somewhat
humid, are covered by vegetation that is unsuitable for cattle forage, and
are inhabited by suslik, great jerboa, steppe mouse, common vole and
mole rat.
Roads. The right-of-way zones along rail and major automotive roads
are characterized by a mammalian assemblage similar to that of the
pastures.
These zones of virgin land are inhabited by little white-toothed shrew,
black hamster [Mesocricetus auratus nigriculus], common
hamster, great jerboa, southern birch mouse, mole vole, mole rat and
little suslik. The weeds and ditches along the railroads are attractive to
long-eared hedgehog, tiger polecat, Siberian polecat, weasel, steppe
mouse, striped field mouse, common vole and migratory hamster.
Cultivated areas. The composition of the mammalian assemblages
in cultivated areas depends upon the routine agricultural techniques, crop
rotation and the size of the arable areas. Usually large areas in cereal
crops are inhabited only by common vole and steppe mouse. The margins
of the planted areas are inhabited by little suslik, hamster, migratory
hamster, mole vole and mole rat (Obolenskii, 1935),
The habitation of the fields by hare is contingent upon the state of the
crop and the method of crop treatment. If rodents and insects are available,
the fox is also a field inhabitant. Thus, for example, when snow cover is
thin in the winter, the European hare feeds on winter crops but rests during
the day in fall-plowed fields. In such a winter foxes will concentrate in
winter-crop and stubble fields which are inhabited by rodents. The most
varied ecological assemblages occur in areas where the plantings of various
crops alternate with sections of virgin land in a mosaic-like pattern.
Perennial crops of alfalfa and other fodder plants are habitats of hamster,
vole, steppe mouse and mole rat. Large plowed fields planted to grain and
regularly worked are characterized by very impoverished assemblages
consisting of fox, European hare, common vole and steppe mouse.
The specific composition of the rodent fauna changes with each year that
land lies fallow. The first year, the stubble may be inhabited by steppe
mouse and common vole, rarely by common and striped field mouse,
526
and even more rarely by birch mouse and migratoryhamster, ''Aging'' of the
fallow land and development of weed cover (green bristlegrass, saltbush,
wormwort) rapidly increases the stability of the rodent populations. The
resulting assemblage consists of long-eared and common hedgehog, fox,
hare, common hamster, mole vole and mole rat. Lands which have been
fallow for many years are usually inhabited by suslik, jerboa and steppe
lemming; this assemblage is similar to the original assemblage.
Fieldcamps, silos and haystacks are inhabited by white-bellied and little
white-toothed shrews, steppe mouse and common vole.
Settlements. The settlements of the Ciscaucasian steppes are
characterized bya fairly large number of biotopes: limestone walls around
estates, ditches overgrown with weeds, gardens, vegetable plots, farm
buildings and stacked sunflower, corn and Italian millet stems (used for
firewood) and invariably inhabited by white-toothed shrew, weasel, Siberian
polecat, house mouse, Norway rat, common hamster and vole. Migratory
hamsters live in non-residential structures. Common field mouse and
forest dormouse inhabit overgrown gardens, particularly those adjoining
exposed rock or groves in valleys and bairaks. Banks of dammed rivers
(reservoirs) are inhabited by water vole. Residences and churches in
settlements are the habitats of common noctules and serotine vespertilio,
and more rarely bicolor vespertilio and sharp-eared mouse-eared bat.
House mouse, either native or introduced, always occurs in residential
buildings in settlements and at field stations. All the large settlements
connected with the Don by rail or automobile roads were inhabited by
Norway rat by the 1930's.
By now the synanthropic assemblage of small mammals in the settlements
of the Ciscaucasian steppes can be considered fully developed, as is the
assemblage of field and house pests (see also Stal'makova, 1935).
The spontaneous development of game hunting in the first half of the
20th century continued to decimate the saiga, corsac fox and, later, the
white [Siberian] polecat populations. Since 1925-27, the hunting of the
so-called ''summer'"' fur species has developed rapidly and affected the
populations of little suslik, common and black hamster and great jerboa
(B. Kuznetsov, 1932).
Large-scale exterminating operations against harmful rodents were
begun in Ciscaucasia early in the 20th century. Pest control of little suslik
on the Ciscaucasian steppes was initiated to protect the peasants' crops.
Later the great epidemiological threat presented by this species was
524 discovered and the boundaries of the endemic plague source area were
drawn.
Organized extermination of suslik and mouse-like rodents has been
carried out in Ciscaucasia only in Soviet time. The areas treated each
year with chloropicrin and poisoned food increased from approximately
100,000 ha in 1923 to 3,000,000 ha in 1935. Tens and hundreds of thousands
of haystacks were surrounded by special ditches, andsilos were gas-treated
(Sviridenko, 1925; Vinogradov and Obolenskii, 1926; Fal'kenshtein, 1933;
B. Kuznetsov, 1932).
The scale of the pest control measures undertaken in the 1920's and
1930's only varied with the fluctuations in population of mouse-like rodents
and the occurrences of epizootics in susliks. The present program is aimed
toward the complete extermination of suslik in the Ciscaucasian steppes.
52%
925
It has not been possible to bring this about by currently available techniques
even with concentration of effort and large expenditures of resources and
labor just within the Ciscaucasian endemic focus.
The operations, however, have resulted in a large decrease in population
and density each year. While there have been no reports of successful
extermination of widely distributed species of mouse-like rodents either,
sudden rises in their populations have been prevented experimentally
following Satunin's (1912a) advice on applying extermination methods in
habitats suitable for survival in those years when they are least abundant,
particularly in the early spring (Naumov, 1946).
Introduction of crop rotation, planting of shelterbelts and trapping
susliks for their pelts has had a much greater effect on the distribution
ranges of susliks and mouse-like rodents than pest control measures.
When fall plowing was introduced in the mid-thirties, it resulted in an
abrupt population decline in mouse-like rodents over large areas. The
plowing of the Salsk steppes and planting of shelterbelts also formed
ecological barriers which prevented the susliks from invading the Kuban
Plain. The continued development of shelterbelts and the construction of
large water reservoirs near Stavropol, Novo-Troitskoe and other centers
will create a more humid climate. Dispersion of common and striped field
mouse and forest dormouse is currently taking place and will continued in
the planted forests connecting the bairak and valley shrubs and forests.
Concominantly, the ranges of steppe and common field mouse, common
vole and hamsters will probably extend eastward into the semidesert zone.
The ranges of small five-toed jerboa, thick-tailed three-toed jerboa, northern
three -toed jerboa, little earth hare and midday gerbil will probably decrease.
The development of hunting will promote the population expansion of European
hare and fox.
The introduction of new species of fur animals onto the Ciscaucasian plains
was associated with the landscapes of the lower Kuban, Terek and Sulak.
Forty-five raccoon dogs (Nyctereutes procyonoides Gray —
Figure 198) were released in 1934 in the tugai forests on the Terek near
Kizlyar. In subsequent years the raccoon dogs became widely distributed
in the tugai and reedlands on the Terek and Sulak, migrating far north
through the reed thickets on the coasts of the Caspian estuaries. The range
of this Terek-based population soon touched the range of the Volga population
which had grown from the introduction of the species along the lower Volga
in 1936 and 1939 (Map 94). Hunting of these animals in the Grozny area
and in northern Dagestan commenced in 1943-44 and the next year the yield
from the lower Terek was nearly 1,000 specimens (Map 7; also see
Lavrov, 1946).
Attempts at introducing some exotic rodents were also fairly successful.
In 1932 the South American nutria (Myocastor coypus Mol.)
was introduced onto the plavni of the lower Kuban in the vicinity of
Grivenskaya (43 specimens) and near Lake Shaitan-Kazak, an oxbow of the
Sulak (22 specimens). The success of this experiment was hampered by
the freezing of the water bodies in severe winters when the nutrias, dying
on the ice of cold and hunger, fell prey to foxes, dogs and predatory birds
(Vereshchagin, 1936, 1947d, 1950a).
Semicontrolled nutria breeding, keeping part of the population in cages
during the cold period, resulted in considerable economic success in these
regions in the 1950's.
528
526
FIGURE 198. Raccoon dog
The introduction of muskrat (Ondatra zibethica L. — Figure 199)
into the reedy sections of the Kuban, Yeya, Terek and Sulak met with more
success.
According to the All-Union Scientific Research Institute of Game
Hunting, muskrats were first released on the Kuban plavni in 1944
and in the reedlands of the lower Terek in 1947. The natural dispersion
of the animals was greatly accelerated in the years that followed by the
artificial dispersion which formed a part of the planned operation
(Map 95).
The development of the muskrat populations of Ciscaucasia will be set
back by the continuation of plavni drainage and the construction of dams
and reservoirs for the regulation of the Kuban, Terek and Sulak rivers.
The relatively slow rate of population increase in muskrats in
Ciscaucasia to date can be accounted for by specific features of the river
regimes (e.g., the heavy June floods which kill the young), and by the
tularemic epizootics of the water rats.
Twenty-three specimens of common raccoon (Procyon lotor L. )
captured in Azerbaidzhan were released in 1950 in the tugai area of the
lower Sulak.
This omnivorous predator found favorable conditions for rapid expansion
of population and range in the relict forests of the lower Sulak and Terek,
where aged trees provide hollows, and frogs and fruit of wild Rosales
provide abundant food. The species also migrates along the Sunzha valley
to the approximate latitude of Achaluki.
It is possible to establish raccoon populations in the tugai forests on the
Kuma in the area of Budennovsk-Georgievsk and in the forest tracts of
Pyatigor'e and along the left tributaries of the Kuban. But the expediency
529
of expanding this predator's range is questionable, since it would be а
threat to poultry farming and to game hunting in the fall and spring
(Vereshchagin, 1953b).
The prospects for enriching the ecological assemblages of mammals on
the Ciscaucasian plains with commercial species depend entirely upon the
expansion or restriction of game hunting. The forest belts can be populated
by roe deer and deer, thereby increasing the population of European hare,
if wolves are exterminated and measures against unlicensed hunting are
improved.
NOES.
vy,
^ hi М
j ah heat
FIGURE 199. Muskrat on feeding ground
SEMIDESERT OF EASTERN TRANSCAUCASIA
The central parts of the Kura-Araks lowlands are mainly covered by
characteristic wormwood and wormwood—saltwort, or, more rarely, by
caper and gramineous groupings. Оп the northern, southern and western
527 margins the semidesert grades into the beardgrass steppe (Grossgeim,
1932). Semidesert and steppe formations are developed in places in the
Gori depression between Mtskheti and Surami.
In the inner part of this zone, gallery forests have survived along the
Kura, Araks, lora and Alazan rivers; also pools grown withreed and bulrush,
and sparse juniper—pistachio forests on the Tertiary hills.
The original Holocene assemblages of mammals in the areas described
were fairly rich in species. The semidesert and steppe were inhabited by
herds of kulan and goitered gazelle. Probably the insectivores and rodents
of the open plains were generally similar to those of the Recent. The
following species occurred in the area: long-eared hedgehog, long-tailed
530
white-toothed shrew, red-tailed Libyan gerbil, steppe vole, William's and
small five-toed jerboa. The tugai thickets and reed—bulrush formations of
estuaries and residual lakes in the Shirvan, Mugan and Mil'skaya steppes
were inhabited by wolf, jackal, striped hyena, lion, tiger, panther, cheetah,
boar, deer and goitered gazelle.
Fluvial lakes of the Araks and Kura and the Caspian bays abounded in
aquatic birds, both migrating and wintering. These habitats were also
occupied by water rat, otter and jungle cat.
The documented record of human influence on the animal assemblage
of this zone dates to the Bronze Age, when settled and nomadic hunters and
cattle herders began to draw heavily on the stock of large animals of the
steppes (Vereshchagin and Burchak-Abramovich, 1948; Vereshchagin,
1949c). The rapid growth of cattle-herding tribes and large settlements and
the development of agriculture and irrigation brought about the next phase
of anthropogenic influence on the faunal complex of the Transcaucasian
steppes.
The agricultural history of the Shirvan, Mil'skaya-Karabakh and Mugan
steppes, which goes back many centuries, has not yet been thoroughly
studied by archaeologists and historians.
The relics of ancient canals, fortresses and giant mounds in the
Mil'skaya and Mugan steppes indicate that vast areas were put to agriculture
and that irrigation techniques had advanced to a high level (Maevskii, 1902a,
1902b). The burial fields and cultural layers under current study at
Mingechaur and Uren-Kala are evidence of large settlements in the center
of the Kura-Araks lowlands in the early first millennium B.C. (Sysoev,
1925; Pakhomov, 1923; Dzhafar-Zade, 1946, and others).
The small irrigation systems constructed on the tributaries of the Kura
and Araks are probably even older. Small communities could readily draw
upon the water of these tributaries, whereas the construction of large
canals could only be carried out by a local population at a higher state of
industrial development. In this respect Latynin's (1935) statements on the
history of irrigation in central Asia are equally applicable to Azerbaidzhan.
The oldest oases in the Kura-Araks lowlands are marked by the following
towns and villages: Barda, Ardash, Geok-Chai, Shil'yan, etc., located on
the Tertera, Aldzhigan-Chai, Geok-Chai, Gerdyman-Chai and Akh-Su.
Cereals and, later, cotton were raised in the semidesert, which also
provided winter forage for the cattle of nomadic herders. But it was the
plowing and irrigation of the semidesert that had the greatest effect on the
528 landscape and animals of the Kura-Araks lowlands. Destruction of the
tugai forests, winter cattle-grazing of the vegetation cover and drainage
of river lakes and flooded areas were of secondary importance to the wild-
mammal fauna.
Preliminary observations on reptiles and amphibians in the area of
newly-established cotton sovkhozes in the Mil'skaya steppe show that
irrigation produces differentiated effects on species populations: it causes
extermination or decrease in some, and growth in others (A. Bogachev,
1938). Sudden changes in the composition and abundance of nesting and
wintering bird assemblages occur when irrigation is installed in semideserts
(Satunin, 1912a; Vereshchagin, 1950b). In the case of the mammalian
fauna, the evolution of its specific composition on irrigated and unirrigated
lands depends upon the season of the year and the agricultural techniques
531
529
employed, 1.е., the type of the main crop, rotation and treatment of crops,
irrigation and development of shrub and tree plantings between the fields.
The contemporary Apsheron Peninsula with its orchards, vegetable
gardens, trade centers and settlements is a good example of qualitative
and quantitative changes in mammalian assemblages caused by the
development of agricultural and industrial settlements in the semidesert.
At present there are 18 species of mammals on the Apsheron, as
contrasted with 15 in the neighboring areas of the Kabristan semidesert.
There are 5 species of small mammals in the cultivated areas of the
Apsheron which do not occur in the adjacent virgin semidesert: Norway
rat, black.rat, house mouse, Kuhl's vespertilio, common hedgehog.
Goitered gazelle and wolf have been exterminated from this area. The
populations ofhare, migratory hamster and steppe mouse increased as the result
of new protective vegetation and fodder plantings (Vereshchagin, 1938b,
1949c).
The concentration and distribution of bats furnishes interesting examples
of the effects of human activity. Bats appear in the semidesert and settle
at structures as isolated as water towers, tombs, sheep pens and, in the
oases, in the hollows of old plane and walnut trees.
Between the years 1935 and 1945 we observed and captured specimens
of the following bats inthe agricultural settlements of the semidesert zone:
Rhinolophus ferrum-equinum V. nathusii
Rh. hipposideros V. pipistrellus
Myotis mystacinus У. serotinus
Vespertilio kuhlii Nyctalus noctula
All these species use residential and service structures for their day
resting places; they feed on the outskirts of settlements and along the
borders of gardens and parks (Vereshchagin, 1942b; Kuzyakin, 1950).
They mainly inhabit crevices under roof beams and openings in the corners
of window frames of railroad stations, schools and other buildings
constructed of cut limestone blocks or bricks, and the stone domes of
mausoleums and mosques. The vespertilio, particularly the Mediterranean
species (Vespertilio kihlii), has become adapted to catching insects
near electric light. In the urban center of Baku, large as it is, У. kuhlii
is now the dominant bat species, althoughin the 1930's it was exceedingly
rare there. We believe that its appearance and increase in numbers over
these decades can be attributed to expanded cultivation, the growth of the
flying insect population and the construction of freshwater pools as watering
places. Following a certain route through the central sections of the city
in 1935, I noted1-3 specimens of vespertilio, in 1940 — 3-5 specimens,
and in the summer of 1949 — 3-7 specimens.
The examples of animals belonging to other orders were selected from
areas irrigated by local canals and consisting of small fields alternating
with virgin land.
Observations made in the Shirvan and Mugan steppes on changes in
composition and relative abundance of the surviving mammals caused by
irrigation and planting are given in Table 107.
332
TABLE 107. Changes in specific composition and relative abundance of mammals occurring with the
irrigation of the eastern Transcaucasian semidesert
Species
Insectivora
rlemiechinus
ВУЗ otoeelio aswanoNs
Crocidura russula
guldenstaedti
© ИСО Sogo о
* Erinaceus euro-=
DACWR Sonoanedo06
Carnivora
Vulpes vulpes ....
GAMLS UM)OUMS Goods 6
MEIGS MGS донос
Vormela peregusna
Mustela nivalis
Ста ао G6 66.0 o
SVP AMG CVE WS Uy a bea. b
Lagomorpha
Lepus europaeus...
Rodentia
Mus musculus
GALERIE Фо рыб .
Cricetulus migra-
О О с Aen ОКО в
Microtus зоста 11$
Meriones eryth-
PO (TWIG: whale ее :
Allactaga williamsi
Ло GWA GET 5 oo 8 bh cols
“Mus musculus
199) 10S CTU ING д оо a geo
* Apodemus sylva-
о met leh ee Ь с
* Вани та tb US ee oy.
2 MCMV Sli CWS 5 66 4
"Dyromys nitedula
Virgin
semidesert
PSS PS 25 OS
Wav iia le
Cotton
Barley
XXXXXX
333
XXXXXXX
Two-year-old
wasteland
x
XXX
Road
shoulders ,
ditches
with weeds
Untended
gardens
in villages
XX
XXXX
TABLE 107 (continued )
Road
Ре ac} Untended
| Virgin [2] shoulders ,
Species к iM р gardens
semidesert a ditches Е Е
Onis : in villages
9 with weeds
=
Е:
НЕ
Artiodactyla
Gazella subgut-
тогоза ..........
Notes. 1. Symbol Х designates presence and relative abundance of species.
2. Data from summer and winter counts are combined for virgin land, waste land, road shoulders and
gardens. The data on the presence and number of animals shown in the fields is drawn from large areas
planted to winter crops. During irrigation almost all the species are driven from the fields.
3. Species migrating from other zones and biotopes are indicated by an asterisk.
4. Table does not include water rat, which occurs only in the Agdam oasis and is absent from other lowland
oases,
5. For widely distributed species — jackal and fox — only the habitats where they feed and rest are given
in the table.
The data show the greatest changes in the original assemblage of
semidesert mammals when the semidesert is planted to cotton. The game
animal population, however, is promoted by inadequate agricultural
techniques, local methods of irrigating with an insufficient water supply;
alternating sections of virgin land and waste land, and abandoned ditches and
plantings of tree and shrub thickets.
The species composition of the animal assemblages is most variable
in old oases with long untended gardens, particularly those adjacent to
tugai and lowland forests. All the new immigrant species which invade
the human environment originate in gallery and lowland forests and reed
Swamps: common hedgehog, jackal, jungle cat, black rat, common field
mouse, forest dormouse and boar.
The mammals which inhabit the vast semidesert areas irrigated by a
system of water canals live under a variety of environmental conditions.
The complete displacement of steppe vole and gerbil from the well-irrigated
lands of the first, second and third Mil'skaya sovkhozes was recorded in
1936-37. White-toothed shrew, hedgehog and steppe mouse, however,
survived on the embankments of the canals, because ground holes are not
so essential to their existence.
The populations of these species were stabilized and steppe vole, hare
and fox appeared in these habitats in later years as weed development and
planted tree stands took hold on the banks of the ditches.
Some specific examples of the reactions of various species to
environmental changes in the semidesert are given below.
534
531
Insectivores and rodents. The regular heavy irrigation of the
cotton fields drives the long-tailed white-toothed shrew, steppe mouse,
gerbil and steppe vole off the land. The environmental conditions created
by weeding the cotton fields are unsuitable for rodents, at least until the
start of winter. The two-stage irrigation method used in grain and alfalfa
fields affect the animals less adversely, as they can find refuge in the
"dry gaps, '' the margins of the fields and the banks of the canals during
irrigating. When the animals reinvade the dry fields, their reproduction
rises rapidly, particularly in the case of the steppe vole (Rodionov, 1924)
which inhabits alfalfa fields in masses. The increase in the rodent
population and in its harmful affects is directly related to haphazard
alternation of planted fields with waste land of different ages and with
virgin land, and to inefficient agricultural techniques (Vereshchagin, 1942c,
1946b).
The irrigation network is very important to the development of viable
assemblages of small mammals in the semidesert. Canals under
constant use are sometimes inhabited by water rat and also serve as
dispersion routes for this species.
On a nutria farm at Karayazy water voles and Norway rats dug holes
in the ditches which collect and distribute water to the concrete cages of
the nutria. At this farm the water rat has become a synanthropic species
which feeds on the residue of grain and roots washed out of the cage basins.
Steppe vole, red-tailed Libyan gerbil and steppe mouse usually inhabit
the outer slopes and tops of banks of irrigation canals, attracted there by
the good aeration of the soil, the ease of hole-digging, the rapid runoff of
rainwater and the growth of weeds suitable for forage.
The remains of ancient irrigation ditches in the steppes, often
recognizable only as small depressions and low ridges, are invariably
inhabited by steppe vole and red-tailed Libyan gerbil. These old ditches,
overgrown with green ephemerals and shrubs of saltwort and caper, serve
as important dispersion routes for the animals during the fall when
population and activity increase.
Railroads are similarly significant to the life and distribution of rodents
in the semidesert. The elevated sections are regularly inhabited by steppe
vole, red-tailed Libyan gerbil, steppe mouse and long-tailed white-toothed
shrew. The soft sandy soil, suitable for hole-digging, the weed vegetation
rich in'srains (Xanthium strumarium) "Cirsium Artie mi,siray,
Salsola, Sueda), andthe seeds of melons and bits of bread thrown out
of the railroad cars — all these attract the gerbils.
The gerbils occasionally dig their holes under the crossties in much
the same way as they dig under stones: they penetrate the soil at the face
of the crosstie and burrow deep into theembankment. These holes are
constantly disturbed by passing trains which leave an accumulation of
debris at their entrances, and by the repacking of the excavated sand by
railroad maintenance crews. Nevertheless, the gerbils persist in
renovating them over and over.
In the summer steppe voles inhabit the lower sections of the shaded
northern slopes of the railroad enbankments, where the soil is cooler and
moister and there are more succulent plants. In the winter they settle
on the southern slopes where the growth of ephemeral grasses begins two
to.three weeks earlier and proceeds faster than on the northern slopes.
S55)
532
The shoulders and ditches beside both unsurfaced and hard-surface roads
are important to the dispersion and maintenance of the micropopulations
of steppe vole. Even in years of decline in population and reproduction,
viable colonies survive in such areas.
The great adaptability of William's jerboa is displayed in its hole-
digging on soft-surface roads in spite of the noise, shocks and clogging of
the holes with debris from passing carts and automobiles.
The populations of the European hare of the Kura area fall into two
different categories, one of which may be called the ''domestic'' population,
the other the ''wild.'' The first inhabits the outskirts of settlements, where
they usually remain by day among weeds and heavy growth of Imperata
cylindrica, often as close as 30-50 m from the yards of houses.
A thorough search near small settlements (e.g., ten to fifteen clay brick
houses) may reveal three to five hares. They are not disturbed by the
usual daytime noises of the village — the movement of people and cattle,
and the barking of dogs. A zone 2-3 km wide, which is completely devoid
of hares, begins 80-100 m from the settlement. Beyond this zone the hares
of the "ма" population begin to appear. It would appear that the ''domestic'
hare habituated itself to life near human settlements because it was
relatively undisturbed and the dogs afforded protection for its day resting
places from foxes and jackals.
For a long time the distribution of rodents in the semidesert and the
viability of their populations has been noticeably affected by the driving of
cattle, the winter-pasturing of large herds of cows, sheep, camels and
horses and the establishment of cattle-herders' stations. In past centuries
these stations consisted of a few felt tents for living and a number of pits,
surrounded by reed screens, for the protection of the young cattle. They
were usually located on ancient alluvial ridges, which are highly attractive
to the red-tailed Libyan gerbil. When the sites were abandoned they
remained for decades as depressions and manured, overgrazed fields
overgrown by milk thistle, white wormwood and orach. These sites are a
characteristic element of the landscape of the Kura-Araks steppes.
It is on these ''wounds of the virgin land'' that the viable populations of
the steppe vole and red-tailed Libyan gerbil usually survive the years of
low reproduction. High weeds protect the voles from dangerous heat
exposure in summer; fast-growing ephemerals provide forage in winter.
The abandoned pits are used by long-eared hedgehogs, white-toothed
shrews, foxes, badgers, tiger polecats and ruddy sheldrakes for digging
burrows and building nests and shelters.
Over the millennia the overgrazing of the wormwood-—grass cover of the
semidesert greatly limited the size of the local populations of vole, gerbil
and jerboa, and affected their redistribution.
In newly-built villages — temporary or permanent — the synanthropic
assemblage of mammals develops at first from local forms: long-tailed
white-toothed shrew, steppe vole, migratory hamster, steppe mouse and
black rat.
In the mud huts of fishermen and hunters built around reed lakes on the
lowland, steppe mice and long-tailed white-toothed shrews are very common
and are a great nuisance.
536
533
Norway rats are always found in profusion near the fisheries оп the
lowland lakes. Steppe voles rapidly invade the new huts and tents of nomadic
cattle breeders in the Shirvan and Mil'skaya steppes. They steal bread
and other food, unperturbed by the presence of people. The migratory hamster
always inhabits residential structures, primarily in the southern part of
the lowland from the village of Pushkino and further south, and in the
foothills of the Lesser Caucasus and the Talysh uplands.
As the settlement continues to develop, cosmopolitan synanthropic
species appear, sometimes after several decades — house mouse, black
rat and Norway rat. The Norway rat, however, also emerges from "мПа"
natural environments (i.e., from populations inhabiting the reed marshes
of Transcaucasia) to invade human dwellings. This is also true of the
black rat originating in local relict colonies.
The author's expedition of March 1940 to the foothills of Karabakh
yielded the following sampling, totaling 1,536 animals trapped in the houses
of the larger villages:
Mus musculus musculus — 78.2%
М. musculus abbotti — 12.1%
M. musculus tataricus — 2.7%
Microtus socialis — 1.4%
Cricetulus migratorius — 1.2%
Crocidura russula guldenstaedti — 0.3%
Tables 108-110 set forth the small-mammal yield from traps and the
large-mammal count in various types of human settlement in the semidesert
zone as examples of the continuous development of synanthropic (house)
and oasis assemblages.
TABLE 108. Species of small mammals from sheep-breeders' winter station in caper formation in the
center of the Shirvan steppe (5 April 1939)
Number of
specimens per
100 traps over all
the sites
Species Trapping site
Insectivora
HAMICCIMIMUS АМИ sod ao oo
Grociaiwmra WMememreloim" io 485 ah15 5.8
Pit overgrown with weeds
Pit for keeping lambs covered with reed roof
Rodentia
Mierlomes (er yar nino ns) sv sis eyes
Mus musiemius: tatarieus er .).
On enbankments near abandoned pits
2 specimens from a tent; 4 specimens
from sheep pen
6 specimens from area between tents;
10 specimens from tents
Мисти оста,
537
TABLE 109. Species of small mammals from the village station of Kerar in the Shirvan steppe
(10-15 April 1939)
Species
Insectivora
Crocidura russula
ри 1 Чепзтае@ т!
ен er)
Chiroptera
oe eee eee та
Vespertilio ktihlii
У. serotinus
ооо
Rodentia
Ми muse uwlis mmiUsicwiit si . es
M. musculus tataricus
© ie) ее ©
Rattus NOTVERICUS. . 6. cs ewe es
Number of
specimens per
100 traps over
all the sites
Trapping site
In firewood storage sheds
In garret of station building
In residential barracks
In firewood storage sheds and poultry
houses
In residential barracks
Note: The village consists of ten wooden residential barracks and a stone station building. There are
a few young acacia trees.
Traps set at the same time in the surrounding semidesert yielded
15 steppe voles and 2 red-tailed Libyan gerbils for 100 trap-nights. In
addition, 2 foxes, 1 hare and 5 goitered gazelles were counted near the
field station.
534
At the time of the sampling, steppe voles and red-tailed Libyan gerbils
inhabited the surrounding open wormwood—saltwort and cereal—
ephemeretum semidesert, as well as the unirrigated experimental barley
fields.
Long-eared hedgehogs, foxes, weasels, hares and goitered
gazelles were also either caught or observed in the vicinity of the
settlement.
Observations taken while crossing an untended garden (area 15 ha) on
the outskirts of the settlement showed 3 hares, 2 foxes, 5 jackals and
1 jungle cat.
In the large cities of the lowland the synanthropic assemblage is
impoverished because of the disappearance of the local rodent and
insectivore species.
Baku is an example of this phenomenon: Kuhl's vespertilio, house
mouse, Norway rat and black rat are the only four species of small
mammals to occur within the city limits.
Carnivores. Wherever fox, jackal, weasel, badger, tiger polecat
and jungle cat appear and prosper in the oases and among the agricultural
535 settlements in the Transcaucasiandesert, arelationship can be established
to the available food resources (insects, rodents and poultry), a favorable
relief and protective vegetation cover, which is particularly important
538
in raising the young. These environmental factors are still dominant
even where game hunting is widespread.
(534)TABLE 110. Species of small mammals from the town of Agdam (24-30 April 1940)
Number of
Е specimens per Е F
Species Trappin te
P 100 traps over aaa as
all the sites
Insectivora
Crocidura russula
ие пзваеч а о... Bacoge . In gardens and alfalfa
Co ПЕСО о боово ов ооо во вое In alfalfa
Chiroptera
Vespertilio pipistrellus....... Killed in garden
ПЕ УЗЕЛ 615 Go a alc ее : In garret of residence
Carnivora
MESECLE ИЕ oo capone о в coo Killed in garden
Lagomorpha
Lepus europaeus cyrensis..... Killed in garden
Rodentia
Mus musculus musculus....... In warehouses and residences
М. musculus tataricus ....... In gardens and stacks
Cricetulus migratorius ....... In gardens and edges of fields
Memos Бек еее ooo ods On sunny empty lots in settlement
Apodemus sylvaticus arianus
In garden ditches
Миегови оса ee ves cite lene
AVG O lia (VETTE SiUTIS и, еее таль
In alfalfa and winter barley crop
In banks of irrigation ditches in gardens
Note, Town in the Karabakh steppe with apple orchards, vineyards and gardens surrounded by barley and
alfalfa fields and old wasteland. Sampling in the alfalfa and barley fields and in the buildings was done
in 400 trap-nights; sampling in gardens, ditches and waste land in 200 trap-nights.
Ungulates. The goitered gazelle and boar provide some interesting
examples of existence and behavior in the cultural landscape of eastern
Transcaucasia. Although the goitered gazelle has been an officially
protected animal during the last 30 years, it is gradually being exterminated
by unlicensed shooting from cars and by shepherds. It is also being driven
from its habitats by destruction of its feeding grounds and formation of a
soft-ground footing by plowing and irrigating the steppes. The species lives
well, however, among the unirrigated cereal crops on the Adzhinour
plateau. The animals remain in the area even when the barley and wheat
are high in May and June, because the soil of the dry fields is sufficiently
firm to allow their escape from wolves. The goitered gazelle is persistent
539
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536
in its habitation of the saltwort—rape thickets covering wasteland 2-3 years
old in the Mil'skaya and Shirvan steppes. This could be particularly well
observed on the lands of the Mil'skaya sovkhozes in 1936 (Vereshchagin,
1938b) (Figure 200).
The first flocks of sheep and herds of cows and camels which commence
to fill the steppe after October cause a redistribution of the goitered gazelle
population. They at first occupy the poorer pastures and salines. Because
of the increase in the number of herds and herding stations, and the flooding
of the salinas in mid-winter, the gazelles are forced to graze near the
the sheep, and sometimes among them. This type of distribution, however,
is always temporary and unstable.
When goitered gazelles are persistently pursued by mounted hunters,
they are likely to change habitats abruptly, hiding for two to three days in
tugai brush and reeds. They flee into the salinas and to hillocks and gulleys
from hunters in automobiles.
As oases were developed, boars took up habitats near the settlements.
Their day resting places are now commonly found among dewberry bushes
in gardens in Agdash, Karabudzhakh and other towns and villages of the
Shirvan and Mugan steppes. Their lairs often occur on the banks of
reservoirs and in ditches overgrown with reeds and plumegrass in the
proximity of residences and traveled roads.
A secondary anthropogenic effect on the mammals of the semidesert was
produced by the destruction of tugai forests on the Kura and Araks and
their tributaries.
In eastern Georgia the tugai forests began to disappear rapidly in the
19th century, until by the mid-century they were presumably placed under
protection and developed for economic purposes through reforestation
(Ketskhoveli, 1942). The tugai forests of Azerbaidzhan were being rapidly
destroyed as early as the first half of the 20th century.
As the tugai vegetation disappeared, bear, panther, tiger, lion and deer
were displaced from the semidesert and the range of boar shrank.
This brief review of the direct and indirect effects of human activity
on the Holocene faunal complex of the Kura-Araks lowlands shows that it
resulted in an impoverishment in large mammals. Some enrichment in
the specific composition of the complex was caused by the appearance of
bats and the enlargement of the ranges of small mammals inhabiting
protected biotopes (white-toothed shrew, common hedgehog, jackal, hare,
common field mouse, black rat, etc.).
The destruction of tugai forests and the expansion of cultivated and
irrigated areas resulted in the gradual enlargement of the ranges of small
semidesert mammals on the margins of the semidesert, and in the
migration of forest species tothe central parts of the semidesert.
The most recent changes in the ecological assemblages and faunal
complex of the semidesert mammals in eastern Transcaucasia were caused
by purposeful human activity: the continuing efforts to exterminate the
steppe vole, red-tailed Libyan and Asia Minor gerbil, jackal, wolf and
jungle cat, and the drawing upon wild-life resources (аз game) suchas hare,
fox and boar. The goitered gazelle remains officially under protection,
but, lacking a steppe preserve and adequate supervision, is gradually
becoming extinct.
541
537
Large-scale extermination of the steppe vole, the chief pest in grain
fields, began early in the 20th century through extensive irrigation and the
spreading of poisoned bait (Satunin, 1912a; Rodionov, 1924; Vereshchagin,
1942c, 1946b).
Not untilthe 1940's were attempts made to exterminate the red-tailed Libyan
gerbil by introducing chloropicrin into its holes and by trapping. This was
done on the Apsheron Peninsula and, to a lesser extent, on the Mugan and
in the Karabakh foothills. In no instance were these operations successful
in reducing the total distribution area of this gerbil, although in years of
high reproduction rates they were carried out in areas of many hundreds
of thousands of hectares. The complete extermination of steppe vole and
gerbil on the Kura-Araks lowlands does not appear to be economically
feasible at the present technological level. With this in mind, it is suggested
that the populations of these pests can be restricted by plowing stubble
fields and using poisoned bait, giving due consideration to the viability of
the populations and weather forecasts (Polyakov, 1950).
The greatest damage to livestock and game is caused by wolves, jackals
and jungle cats, which are far from being completely exterminated by the
usual hunting methods. The statistics of pelt yields of these species for
the years 1925-1955 show that fluctuations in the yields were controlled by
the activity of the fur trade and the amount of the premium paid for each
pest that was killed. For example, a 500% increase in the price paid for
a jackal in 1949 increased the yield by 500%.
Th
НХ (р 2
iW ei
РА Pes 4
er ae qt
ss
a
SN
t
‘aes
os
FIGURE 201. Rabbits grown wild on Bulla Island
Planned introduction of alien species into the ecological assemblages of
the semidesert was carried out in order to increase the yield of the valuable
furs.
In the second half of the last century, rabbits of the species Lepus
cuniculus Г. were introduced by sailboat captains and lighthouse keepers
onto the Caspian islands of Zhiloi, Nargen, Bulla, Oblivnoi and Svinoi. They
542
became acclimatized and grew wild. In 1931-1952 [sic] small groups of
the Viennese blue rabbit were released on the islands of Urunos, Zhiloi,
Bulla, Los! and Oblivnoi in order to establish a rabbit trade.
These varicolored wild animals, whose average weight is 1.5 kg, live
and reproduce on these islands almost entirely without peril from
terrestrial enemies, and unhindered by lack of fresh water (Figure 201).
In 1940 approximately 500 rabbits inhabited Bulla Island (Vereshchagin,
1942b). The only check on their population increase is unlimited shooting
538 by lighthouse keepers and visiting hunters and inadequate food and shelter.
The compacted volcanic mud of the islands is easily eroded by winter rains
and becomes unsuitable for digging holes. The rare Salsola
dendroides, Frankenia, Ephedra, glasswort and shoots of hare
barley and meadow grass, which die out by June, do not provide sufficient
forage. Constructing artificial shelters and supplying additional food would
be the important steps in organizing rabbit farming in this area.
Since 1931 nutrias have been introduced into a number of plain water
bodies overgrown with reedmace—cattail (Figures 202, 203; Map 95).
FIGURE 202. Nutrias swimming and feeding
Photograph by author, 1948
The factors limiting the increase and dispersion of nutria populations
were frost, winter-freezing and summer-drying of water bodies. Under
these circumstances large numbers of nutria are easy prey for jackals
and dogs (Vereshchagin, 1936, 1939c, 1942b, 1950a). The natural
543
dispersion and range development of nutria followed the basins of the
interconnected lakes, Shil'yan and Kara-Su, at the rate of 15-20 km a year.
The populations and the yields, however, were unstable (Graph 20).
At present stable natural populations of nutria occur only on sections
of spring-fed rivers which never freeze, e.g., Lake Khuluflu in the
Shamkhor District and the Kara-Su rivulets at Karayazy. Notwithstanding
these difficulties, there are good prospects for the free breeding of nutria
in Azerbaidzhan if certain conditions are met: a regular water supply to
the basins, the availability of winter corrals and a stock of animals in the
corrals for emergency situations, such as heavy frost or drought.
It can be anticipated that large-scale irrigation operations and
afforestation will produce much more pronounced changes in the semidesert
ecological assemblages in the near future. The construction of the
Mingechaur reservoir has already created the possibility of irrigating
another million hectares of steppe land.
FIGURE 203, Cattail thicket, thinned by nutrias, on Lake Shil'yan
Photograph by author, 1948
The construction of the irrigation network will allow heavy repeated
irrigation, further expansion of cotton planting and grassland crop rotation.
Improved methods of extermination used on the dry gaps, canal banks and
other places will depress the populations of jerboas, gerbils and steppe
voles in the vast lowland areas. The planting of shelterbelts and shrub
thickets will expand the distribution ranges of white-toothed shrew, common
hedgehog, jackal, weasel, jungle cat, hare and steppe mouse and, toa
certain extent, black rat.
544
540
Hunting, therefore, must be controlled and measures taken to prevent
the development of dangerous epidemiological conditions.
Any further enrichment of the mammalian complexes of the
Transcaucasian steppes and semideserts by new species of game animals
has limited prospects in the environment of the cultural landscape.
Commercially useful rodents, as well as those that are pests, are not
tolerated in fields of grain and industrial crops. And, lacking the rodents
which are their chief food supply, small species of furred carnivores either
die out or begin to prey on poultry and wild birds. Therefore, the project
of introducing Siberian polecat into eastern Ciscaucasia, where the rodent
populations are subject to considerable fluctuation, is economically and
biologically unsound. Wild ungulates such as goitered gazelle, saiga and
some other African antelopes (gnu, bubaline and other species) will be
able to live on those sections of land which are unsuitable for irrigation
and agricultural development, e.g., in Kabristan and the Dzheiran-Cheli
steppe.
The Pleistocene occurrence of saiga in Transcaucasia and the experimental
release of 20 specimens on Bulla Island in 1950 indicate that the
introduction of this species is quite possible. Clearly, however, without
adequate protection, these new immigrants to the steppes would be as
disadvantaged as the goitered gazelle.
FOREST-STEPPE AND FOREST ZONES
In the development of the most recent ecological assemblages and
mammalian complexes of the forest-steppes and forests of the Caucasus,
the anthropogenic factors are interwoven in a complicated pattern with
the natural factors.
The postglacial mountain landscapes of the Caucasus formed under
conditions of general warming and lower humidity. As the glaciers melted,
vast areas in the glacial valleys and on the slopes of the Main
Range in the western Caucasus became covered by meadow and mountain-
forest vegetation which migrated from the lower zones and over the spurs
of the longitudinal ranges.
Biocenotic relationships of the Recent type formed in the mountain-
forest zone as developing forest groupings expanded over the subalpine
meadows (Tumadzhanov, 1947). The continued warming and increasing
drought created an encroachment of steppe formations on the forests.
The uplift of some sections of the foothills forest-steppe zone during
the xerothermal phase of the Holocene was considerable and, because of it,
chernozems developed at altitudes as high as 1,000 m on the western spurs
of the Greater Caucasus and near El'brus (Yakovlev, 1914; Zakharov,
1935; Berg, 1947; Klopotovskii, 1948).
During the next phase of the Holocene (the beginning of our epoch), the
process probably operated in reverse with the forest formations
encroaching on the steppes, a development which was particularly
pronounced on the piedmont plains.
The cenotic evolution of Caucasian forest assemblages in the Quaternary
has been discussed by Sinskaya (1933) and Kolakovskii (1947a) on the basis
of geobotanical analysis. In the present epoch, however, the natural
545
541
development of successive formations of trees was not significant in the
formation of landscape and ecological assemblages of mammals; indeed,
haphazard cutting and burning of forests and cattle grazing greatly
outweighed the natural processes. Because of human activity the oak,
pistachio and hornbeam forests of the foothills degenerate to thickets of
xerophytic deciduous thorny shrubs, which are later replaced by beard-
grass steppe, or even by phrygana. Cutting of the mountain beech forests
promotes the growth of hornbeams, and grazing further results in the
development of birch groves and rhododendron thickets, which finally grade
into subalpine meadows (Bush, 1935; Yaroshenko, 1940, 1945).
According to botanists and soil scientists, the foothill and plateau
forests of the Caucasus covered a larger area in the earlier historical epoch
than they do at present. Their conclusion is based principally on the
occurrence of relics of ancient forests in areas which are now steppes,
particularly in Ciscaucasia and Armenia (Bush, 1900, 1935; Kuznetsov,
1900; Troitskii, 1927, 1939; Takhtadzhyan, 1941; Grossgeim, 1948).
The formation of other plant assemblages in areas of destroyed forest
and new forest growth in various regions of the Caucasus was undoubtedly
controlled by specific local conditions created by the ecological situation
and the natural process of vegetation replacement.
However, the most recent natural trend caused by climatic changes
toward replacement of steppe formations by forests (Zakharov, 1935) and
conversely of forests by steppe formations (Yaroshenko. 1940, 1956) is
completely overbalanced by human activity which always leads to the
development of xerophytic vegetation (Map 97).
Man's early influence on the mountain forests was mainly reflected in
the contraction of the upper and lower boundaries. But commercial
deforestation during the last centuries has exposed the slopes on a large
scale and caused the formation of forest glades and woodless belts over
the entire altitudinal range (see Zaklinskii, 1931; Solntsev, 1940).
Deforestation in the Caucasus produced these general consequences:
the soil became desiccated; vegetation of a definitely xerophytic type
replaced the forests; the climate changed, becoming less humid and having
less atmospheric precipitation, because fog no longer condensed on the
leaves and branches of trees; erosion increased sharply and as the soil
layer was removed, gulleys and mudflows formed (Kirillov, 1915; Roshchin,
1928, 1930, 1931).
Almost everywhere the destruction of the forests brought about a
displacement oftypical forest ungulates — deer, roedeer, elk, bison; of forest
carnivores — bear, pine marten; andof arboreal rodents — fat dormouse,
Persian squirrel and, partially, Caucasian yellow-spotted mouse. The
effects of deforestation on those rodent species which do not depend heavily
on forest habitats (e. g., vole and common field mouse) were more complex.
Another result was the invasion of the mountain slopes, exposed and
desiccated by deforestation, by relatively xerophilous rodents and
carnivores from adjacent zones of open landscape. Specific local changes
also occurred in each geographic district; these were associated with the
particular composition of the mammalian fauna and the particular structure
of the environment. They are discussed below according to natural zones.
546
542
Ciscaucasian forest-steppe
The broad belt of the Ciscaucasian sloping plains extends from the Taman
Peninsula in the west to the Caspian coast and Mount Beshbarmak in the
east. Its southern boundary is formed by the steep slopes of the Greater
Caucasus, its northern by the latitudinal sections of the Kuban and Terek
and the northern slopes of the Stavropol Plateau.
In the xerothermal postglacial epoch this zone was probably covered by
steppes and in the sub-Atlantic epoch by oak forest alternating with meadow
and meadow steppe on the divides. The contemporary forest-steppe
landscape of this zone is clearly secondary and anthropogenic (Bush, 1900;
Tumadzhanov, 1947, and others).
-The ancient sections of the dry steppe, Lower Quaternary in origin,
have survived only between the Tertiary hills of the Terek and Sunzha
ranges and on the southern cliffs of the limestone ridges of the foothills,
and in the northern foothills of Dagestan.
The archaeological studies of Gerts (1870), Farmakovskii (1914),
Uvarova (1900), Gorodtsov (1935, 1936), Anfimov (1935), Iessen (1940),
Krupnov (1946, 1947, 1948, 1949) and Kruglov (1946a) show Early and
Late Bronze Age settlements built on these piedmont plains. The settlements
are representative of Maikop, north Caucasian, Koban, Scytho-Sarmatian
and Hellenic cultures. In the present era the region was settled by Alanis
who were gradually driven out by migratory Moslem tribes. Mischenko
(1923, 1928) and Zakharov (1935) report that the relict oak—hornbeam
forests of the Trans-Kuban Plain have in this era overgrown the Scythian
mounds which were originally built on the steppe.
There were still many isolated forest islands and ritual groves of the
Adygeians (''kodoshi"' or ''kudoshi'') remaining on the plain in the middle
of the last century, indicating that in the past the area was covered by
extensive tracts of forest (Veidenbaum, 1878).
The destruction of these forests by the local population was nearly
complete by the time of the first large-scale settlements of the Cossacks
in the mid-19th century (Felitsin, 1884; Shcherbina, 1913).
Their large settlements of warriors and hunters were built at that time
in the valleys of the left tributaries of the Kuban and the right tributaries
of the Terek and Sunzha. These valleys grown with tugai thickets were
main routes for the seasonal migrations of large animals (boar, roe deer,
deer, bison) fromthe mountains tothe plain, and were particularly important for
the survival of these species during catastrophic snowfalls in the mountains.
The deforestation of the divides and the establishment of villages (''check-
posts'') in the valleys therefore disrupted the established pattern of
migration and led to death from starvation which, along with hunting,
brought about the extermination of the ungulates. The boar, forced to
remain in the mountain forests during winters of heavy snow, was
particularly affected. The clearing of tugai thickets in the lower floodplain
valleys of the left tributaries of the Kuban and the concomitant changes in
the hydrological regime of the streams resulted in the disappearance of
river beaver from this area.
The continuing replacement of forest-steppe by the contemporary
cultural landscape on the Trans-Kuban Plain with great areas of arable land,
pastures and settlements caused the displacement of pine vole to the
547
543
foothills, reduction in the population size and distribution area of the
common field mouse, and an increase in the population size of the
common vole, Asia Minor and common hamsters, migratory hamster,
steppe mouse, mole rat, hare and fox.
Overgrazing by cattle formed bare dry areas on the bottoms of the
valleys and near the canyon mouths; this promoted the rapid migration
of the species mentioned to the zone of the mountain forests, as, for
example, in the vicinity of Krymskaya, Il'skaya and Goryachii Klyuch.
On the sloping Kabarda Plain small forest islands were preserved only
in some gulleys and river valleys, and in the Pyatigor'e area.
The width of the deforested zone between Pyatigorsk and Georgievsk
and between Nal'chik and Prokhladnaya is at present 30—40 km.
Only 200 years ago this plain was still inhabited by roe deer, deer, elk
and bison; at present it is characterized by a relatively impoverished
assemblage of synanthropic species and small agricultural pests. The
continued extermination of the forests on the low ridges of the piedmont in
the Nal'chik District results in the drying of the rivers. The mountain
slopes, desiccated by deforestation and overgrazing, remain for some
time covered with nut trees and hornbeams, which gradually retreat up
the slope. Hare, common and black hamsters, steppe mouse, mole rat
and northern mole vole follow this retreat.
Forest belts have recently been planted on the Kuban-Kuma divide to
lessen gulley erosion and summer drought.
Similar changes have taken place in the ancient landscapes of the more
humid Terek-Sunzha valley. Near Alagir there is a grove of 300-year-old
lindens, beeches and filberts called Khetag, sacred to the Ossetians, which
was visited by Pallas. The existence of this grove indicates that the area
was covered in the past with well-developed piedmont forests, which were
inhabited by bison, elk, deer and roe deer.
The consequences of deforestation can be seen in the changes in specific
composition and distribution ranges of the mouse-like rodent fauna.
Pine vole and common field mouse have been displaced to the wooded
foothills from the plains, which were then occupied by wheat-crop pests,
common vole, hamster and migratory hamster. In this region the
extensive fields of vegetables and corn and the rivers which never freeze
provide a suitable habitat for Norway rat which does damage to the fields
and vegetable gardens (Bogdanov, 1929). The xerophilous tiger polecat
migrated here from the direction of the arid Sunzha Range; its distribution
area extends to the wooded slopes of the Chernye Gory Mountains (Turov,
1926Ъ).
As the result of cutting and overgrazing, the oak and beech forests of
the Chechen foothills and northwestern Dagestan between the lower Sunzha
and Sulak canyons were replaced by a peculiar bushy thicket consisting
of filbert (Coryllus avellana), pear (Pyrus caucasica), plum
(Prunus divaricata), common buckthorn (Rhamnus cathartica),
Cornelian cherry (Cornus mas) and other species (Grossgeim, 1948).
Of the earlier assemblage of forest mammals in this area only mole,
European wildcat, lynx, forest dormouse, boar and roe deer still occur
in some places; bear, fat dormouse and deer have disappeared.
Between Makhachkala and Derbent some parts of the low foothill ridges
have been planted for centuries to grapes and corn. Of the old forests of
these foothills there now remains a xerophytic formation in which Christ's-
548
thorn (Paliurus spina christi) and hornbeam forests predominate.
All of these changes were favorable for the widespread infiltration of little
suslik, steppe vole and migratory hamster into the foothills. Geptner and
Formozoy (1941) noted that the species which has invaded Dagestan the most
extensively is the vole. However, it can be deduced from the situation in
eastern Transcaucasia that the more mobile and widely distributed migratory
hamster, although less numerous, was more rapidly dispersed through the
deciduous shrub formation in the gulleys.
Farther south along the lower Samur and other rivers on the Khachmas
lowlands, the forests are partially preserved. The continuous wooded tracts
of the past are now reduced to a forest ''network'' — narrow belts, covered
by liana, surrounding large clearings now converted to the cultivation of
corn and vegetables. Even in its present state, however, the region is still
inhabited by some boars, roe deer and numerous fat and forest dormice
and steppe mice.
The steppe vole has migrated from the dry hills of the Kusary Plain to
the deforested sections and forest glades. On the rice fields peculiar
hydrophilous populations of steppe mouse, Mus musculus
tataricus, similarto M.musculus hortulanus, have developed.
Their hydrophilous habits were, of course, formed long ago, since they
544 also inhabit the margins of cattail marshes and are not solely dependent
on the rice fields. These do, however, provide them with abundant food,
shelter and storage places in the demarcation embankments (Vereshchagin
and Mamedov, 1946).
Corn and sunflower crops in melon cultures attract equally large
numbers of steppe mice to clearings as to the old rice soil (chaltyk). This
is due to rapid development of wheat strains: green bristlegrass —
Setaria viridis, reed canarygrass —Phalaris arundinacea,
barnyardgrass —Echinochloa crus-galli. European hare and
common field mice feed in the clearings at night.
TABLE 111. Composition and relative abundance of small mammals (number of animals per 100 traps) in
various biotopes in the Samur forests in October 1945
Edges of
Old forest | forest with Corn and
on liana Богаег-| sunflower ой ant Rice fields
dry lands | ing streams fields АТО (al
and fields weeds
Land under
Total number of specimens
specimens
number of
specimens
number of
specimens
number of
specimens
number of
specimens
w
[)
+
oO
‚а
Я
=
я
Crocidura russula gulden-
ЕО о
Mus musculus formosovi..
M. musculus tataricus ...
Apodemus sylvaticus.....
А AUREUS вы о ее OOo OO
Миепови вос та BSAs 66s
DyrOwesys паи совооос
Gillis 5 с забовоовоное
549
545
Table 111 sets forth the count of insectivores and rodents from trappings
in this district in October 1945.
As a result of the conversion of the Khachmas lowland forests to crop
areas, the ranges of hare, fox and jackal expanded, steppe vole appeared
in the area and the striped field mouse, which concentrates in the margins
of forests, increased in population.
The broad Kusary (or Kuba) sloping plain, covered with Quaternary
gravels and loams, lies between the Khachmas lowland and the ranges of
southeastern Dagestan. The forests of this plain were cut down during the
last centuries and the deforested areas grown over mainly with Christ's-
thorn, hawthorn, blackberry, hornbeam and filbert, or planted to wheat
and barley. Large apple and quince orchards were planted near the villages.
During the present era far-reaching and complicated changes took place
in the distribution and populations of the carnivores, hares, mouse-like
rodents and ungulates of this plain. Bear disappeared; fox, jackal, badger
and hare populations have probably increased somewhat, and steppe vole
and steppe mouse became widely distributed.
The distribution of the common field mouse assumed a moSaic pattern,
corresponding to relict forests and shrub thickets. Abandoned plowed lands
and pastures were extensively occupied by steppe vole and mouse and the
lower boundary of the common vole range moved upward somewhat.
The recent construction of the Samur-Divichi canal and cultivation of rice
favored the formation of hydrophilous populations of steppe mouse.
Fat dormouse occurs in only a few parts of the relict aged forest,
although forest dormouse is still widely distributed.
The prospects for introducing new species into the mammalian fauna of
the Ciscaucasian foothill forest-steppe plains is rather poor.
Desmans have been released at a number of lake and reservoir locations.
Raccoon dog and common raccoon occur in some parts of the Trans-Kuban
and Terek-Sunzha plains (Map 94).
The-immediate future for this zone is further development of gardening,
acceleration of agriculture, rapid population growth and expansion of
settlements. Forest belts should be established in the driest areas. Game
control measures should take into account the moderate increase in the
population of European hare, the increased abundance of fox andthe localized
increases of mink and Siberian polecat. Agricultural planning should be
directed toward the extermination of hamsters and voles in the western half
of the zone, and of steppe voles and mice in the eastern half.
No nature preserves have been established in the area, and by now such
measures are long overdue.
Changes in the landscape and the mammalian assemblages, similar to
those described above, also occurred on the piedmont plains of eastern
Transcaucasia.
Transcaucasian forest lowlands
In the Holocene the warm, humid lowlands of western and eastern
Transcaucasia (Colchis, Alazan-Avtaran and Lenkoran) were covered by
marshland forests with broadleaf species and thorny liana predominating.
550
The nature of these primitive virgin forests has not been adequately
studied (Al'bov, 1892; Sinskaya, 1933; Maleev, 1938; Grossgeim, 1948).
Typical species are Quercus longipes, hornbeam, elm, Pterocaria
caucaSsica, black alder and, in the Lenkoran lowland, Parrotia
persica.
Man's selective destruction of the lowland forests produced many
marginal sections overgrown with lianas: Smilax excelsa, Hedera
helix, Clematis vitalba, Periploca graeca, and glades
overgrown with bracken, blackberry or Christ's-thorn and spiny Juncus.
The mammalian assemblages developed in different ways in various
sections in response to human activity because, while the landscapes are
homogeneous, the topography is diversified,
The relics of human culture have been more completely studied in
western Transcaucasia where monuments of the Bronze Age (Colchidian,
Hellenic and later) are common in Middle and Upper Paleolithic sites
throughout peripheral areas of Colchis (Rion lowlands).
In the present era large settlements with corn plantations and fruit
orchards were established in Colchis. The populations and specific
546 composition of the mammalian fauna were heavily damaged through direct
extermination efforts and shrinkage of the forest-covered areas. In this
region river beaver became extinct at the end of the 19th century, red deer
disappeared later, and boar and roe deer survived only in some isolated
marshes.
In 1930 operations to drain the Rion marshland (Figure 204) by means of
silting and canals were commenced (Zunturidi, 1931; Shaposhnikov and
Vereshchagin, 1932). At the same time the alder forests were cut and the
land was planted to tea, corn, tangerines and eucalyptus.
FIGURE 204, Alder—water lily marshes near Poti —typical habitat of nutria and Norway rat
Photograph by author, 1931
554
All these processes created abrupt changes in populations and composition
of the mammalian fauna.
The populations of jackal and wolf were supported by poultry- and cattle-
breeding activities. Gardens and many marginal sections of relict forests
overgrown with liana promoted an increase in the fat and forest dormouse
populations, while the deforested, arid meadows became inhabited by
pine vole,
The evolution of mammalian assemblages promoted by the drainage of
the alder marshes and other kinds of production-oriented human activity is
shown in Table 112. Our observations, trappings and tabulations were
made in the fall and winter of 1938-39 and 1940 near the eastern shore of
Lake Paleostom and near Lake Bebesyri.
Most of the mammalian species disappeared from areas planted to tea,
tangerine, lemon and tung-tree because the plantations did not furnish the
animals with shelter and food.
The synanthropic assemblage in the agricultural settlements of Colchis
developed partly from introduced species (Norway rat and house mouse)
and partly from native species (black rat and common field mouse).
Shidlovskii (1947) tabulated the following numbers of rodents caught per
1,000 traps in residential buildings on the Black Sea coast: house mouse —
547 158; Norway rat — 125; black rat — 17; common field mouse — 13. The
latter species does not occur in the larger settlements of the lowland.
Stepanov's data on rodents caught and studied by the anti-plague laboratory
at Batumi show for an 18-year period: 44,771 mice, 40,000 Norway rats,
3,505 black rats and 1,588 Alexandrine rats. According to Stepanov, the
mice and rats are approximately equal in numbers at Batumi. * The mice
mainly inhabit residential (apartment) buildings, while rats live and feed
outside the apartments.
548 Clearly the general trend of evolution of the mammalian assemblages
during the development of the contemporary cultural landscape of the
Colchis lowlands is toward extreme impoverishment of the assemblages.
The large ungulates and medium-sized carnivores disappear, and the
populations and species composition of most local rodents decline.
Human activity in the Alazan-Avtaran lowland in northwestern
Azerbaidzhan, which has a different topography, affected the mammals
in a very different way than in the Colchis lowland. The Alazan-Avtaran
lowland is a valley 20-25km wide and almost 300 km long, extending from
the village of Akhmeti in Georgia to the village of Ismailly in Azerbaidzhan.
Relict forests of Quercus longipes and hornbeam remain mainly in
the eastern part of the valley. Human activity here reaches back as far
as it does in Colchis. Remains of settlements of the first millennium В. С.
are known from the vicinity of Belokany, Kutkashen and Nidzh. The
lowland is now quite densely populated, the chief centers being inthe middle
and at the canyon mouths. The mountain rivers have been diverted by means
of additional channels and ditches for irrigation of the rice (Figure 205),
wheat and tobacco fields and the fruit and nut orchards. Man worked the
greatest changes in the landscape and in the mammals of the area by cutting
the forests and subsequently planting walnut, hazel nut, chestnut, apple,
pear, wild cherry, plum, medlar, quince and grape and by plowing the forest
glades for cultivation of barley, wheat, corn, rice, tobacco and sesame.
* This is highly questionable; as a rule, mice are more abundant.
552
549 The margins of fields, banks of irrigation ditches and sections of pasture
became overgrown with hawthorn, blackberry, blackthorn, Christ's-thorn,
plume grass and reeds. The edges of forests and settlements are
characterized by strongly developed annual weeds, shrubs and trees,
particularly European elder and ailanthus.
(547) TABLE 112. Changes in the specific composition and relative abundance of mammals caused by drainage
and cutting of lowland forests in western Transcaucasia
Forest glades
Forests of with bracken Е
Tea and citrus
plantations
Species Colchidian and marshy Corn fields
type meadows and
pastures
Insectivora
SOE» аа оосовевовосвь
Crocidura russula monacha ..
Talpa ©о08$& с оооовообовосо
Расе GUur@OPBSWS s5o0500c
ES BS <
Carnivora
CAMIMIG ADRES 5 оз особо обю вов
С. ПЕРИ соозобоовоосовов об
Wilpes Уи: совороввоонос
IMIS He ea И МАЙ ее ее аи
MGLOS оао ооо ово номе
Lagomorpha
LEPUS ЭР o5 5660000040
Rodentia
MMS ИБО so 000500506000
ОЕ ии SWUVAEMCWS5S 6505005
RANG HENEEMSS ooo oop odoDGo0000
В ПОРУ ИОВ 5 Sincocagddon6
ПВО МАЕ ОИ 6 оооообвасос
ATWVCOLA воле о Зи 55565 5006
Руза ИЕ S55 noo 5504
Gis о 6-54 oko ne oOo boos
< Муса зоне ур 6 a6 55006
Artiodactyla
SiS wks CHVOID AN 3 st enna он
ооо ооо
Note. Symbol Х designates the presence and relative abundance of each species; asterisk (*) denotes
species introduced either accidentally or purposefully.
The original thickets were often restored on the sites of abandoned
settlements. However, the original species of trees were replaced by
953
different groupings with an admixture of introduced species. For example,
the site of the town of Chukhur-Kabala was overgrown with alder,
Pterocarya caucasica, poplar, maple and species which made a
secondary return in a wild state, such as hazel nut, walnut and plane.
The cutting and burning of the jungle drove out the large animals such
as boar, deer, roe deer, bear, lynx, European wildcat and panther, as
well as pine marten, fat and forest dormice and squirrel, while increasing
at the same time the populations of fox, jungle cat, badger, hare, vole
(steppe, water and common) and black rat.
(548)
> PF é
%
FIGURE 205. Flooded rice field in the Agri-Chai valley — ап example of complete seasonal displacement
of mammals
Photograph by author, 1950
As the nut and fruit orchards age, the assemblage of forest mammals is
almost completely restored. Some species increase because of the
abundant food supply and available shelters in rock walls. These are fat
and forest dormice, Persian squirrel, common field mouse (to some
extent), Transcaucasian subspecies of the Caucasian yellow-spotted mouse,
mountain house mouse and long-tailed white-toothed shrew. The common
field mouse became a synanthropic house rodent in this region. The changes
in the specific composition and relative abundance of mammals in diverse
habitats of this lowland are shown in Table 113.
In summary, then, the deforestation and planting of the lowlands to
cereal and industrial crops in recent times resulted in the disappearance
of 6 species of game animals (deer, roe deer, bear, lynx, European
wildcat and squirrel) and the increase of 2 species (fox and hare). These
processes also brought about the migration of William's jerboa, steppe
554
(550) TABLE 113. Changes in specific composition and relative abundance of mammals caused by cultivation of
field and garden crops in the Alazan- Avtaran lowland
Forest glades with
Lowland virgin Old nut and
Species cereal crops i
P forest P fruit orchards
and shrubs
Insectivora
Crocidura russula gilden-
staedti PAVERS ROT oe Bie. cent ci ххх хххх
Grylie WicoldsOl 42) swans <. Syeie ОВО ово = —
SOR Ar EMS я ое bo Ooo x x
тип асет ентораей 55 955 Ба x XX
Carnivora
сима ет о tats te! weet elle x хх
С. ПИР ons бебыьзоюооваесо x x
WW DSS Урень. 4A po Blog x x
WRIST ENC OR Basis a SG Gabo о x x
IMAGES URE See ее x x
Маме По не еее x x
ICM eSNG Хе ее еее а ое x хх
ЕВ IWERA cho op ola ads в ae Xx Х (on rivers) x
BVeVIMSW CHAU Sis К ote Юга x хх x
ВЕ SIT SEAN, Reese teste Mckee uate x x x
Ie 14 co бо ше oO Sac 5,58 510 x = x
SF POE W/O MOC SENSIS aa 9 6 boo x = хх
Lagomorpha
Lepus €uropaeus........... = Xx
Rodentia
Mus musculus abbotti...... хх ххх
IN ине но ата цеце, 6 44 = —
Ароетаиз ума те ие. ххх XXXX
АИТ У рее ое ее хх хх
вавво ив asDO MUSH. ое a) olka. = X (mear settlements)
PVC OS ay ravactuld Srayeneucileiisi cl eis = XX (near —
mountain slopes)
SRPIVEMS Oe alas te ANS MP eres, ors — XXXX =
АИ ЩЕУ ОНИ о боев с = Х (on rivers) =
” А Vacitagia wa Wiliams м. . 1. = x —
Dy PONS mice Soda eas 66a x x XXX
(GUNS GUS. опрос зоо cham fea хх = XXXX
ОИ A MOMMA As gone 65.0 x хх
Artiodactyla
ShUS, SIC OM ALS econ ее с хх x
GON MSS CAEN INS SS sents ele сое x oe
Gapreolms capreolws о. x is
Note. Symbol X designates the presence and relative abundance of each species; asterisk (*) denotes
acclimatized species and immigrant species from other zones.
555
551
mouse and steppe vole from the southern steppes and of common vole from
the mountain-forest zone.
In large nut and fruit orchards the assemblage of game species is almost
completely restored, and the number of arboreal rodents is on the increase,
This is, however, mainly a seasonal phenomenon controlled by the fall
migratory pattern which brings these animals, inhabitants of mountain
forests and wooded gulleys, to the orchards which are rich in food.
The synanthropic assemblage of the large market-gardening villages in
the lowland consists almost entirely of local species: common field mouse,
house mouse (variety with dark abdomen), black rat, and, to a lesser extent,
forest dormouse and long-tailed white-toothed shrew.
The garrets and crevices in houses are inhabited by sharp-eared mouse-
eared bat and small vespertilio. The introduced synanthropic forms, house
mouse and Norway rat, are now foundinalllarge settlements interconnected
by highways.
Much more pronounced changes in landscape and fauna occurred in the
lowland-forest zone on the northeastern border of the Karabakh, onthe lower
Tertera and in southeastern Mugan along the northern margin of Talysh,
where the cultural has completely obliterated the original landscape. The
nature of the original features can only be deduced now by inference.
Because the nut and fruit orchards are limited, there is no possibility of
even a partial restoration of the earlier assemblage of forest animals.
Only the occurrences of porcupine in the recent past and of black and
Norway rat in isolated gardens in the present indicate that the region was
previously inhabited by assemblages of forest mammals.
house mouse
a SF
ГПШПУИИШКЕ X M XM
Number of rodents caught per 100 traps
4 1941
3 A.
/ = Norway rat
eon,
2 Pred Praise es
11: black rat
i it i! WV Wi VV хим
FIGURE 206. Variations in abundance of house rodents
(house mouse, Norway and black rats) in Lenkoran and
adjacent settlements in 1941-42
556
552
Most of the Lenkoran lowland has been subjected until now to changes
similar to those described for the Alazan-Avtaran valley. Originally it was
completely wooded with thick Hyrcanian forests of oak, alder and Persian
ironwood, covered by thorny liana. Large forest glades were first converted
to rice fields; later, extensive areas were cleared for tea and tangerine
plantations. Combined with the growth of settlements and the development
of hunting these landscape modifications have reduced the populations of
boar and porcupine and completely driven out bear, tiger, deer and roe
deer. Hares migrated to this region from the Mugan following routes along
deforested sections and tea plantations. The steppe vole, however, could
not migrate either from the upland steppes or from Mugan because of the
wet ground in winter. The common field mouse is completely absent and
hydrophilous populations of steppe mouse, so characteristic of the rice fields
of the Khachmas lowland and southeastern Karabakh, are nowhere to be
found in this region.
The populations of jackal, jungle cat, fat dormouse, black and Norway
rats and house mouse thrived as landscapes developed: large settlements,
orchards and fruit gardens, rice fields, clearings, many strips of dense
marginal forest, thorny thickets and hedges surrounding fields, and
water reservoirs overgrown with reeds. Boar, which feeds on cereal
crops, also remained in the area. There is no large-scale hunting to
decelerate the increase in jackal and jungle cat populations which partly
subsist by preying on poultry.
Since there are fewer aged trees, the fat dormouse often uses the
garrets of residential and industrial buildings as day resting and hibernation
places. Garrets and crevices in wooden and stone buildings are also
inhabited by Myotis mystacinus. The synanthropic assemblage of
house pests consists of only 3 species of rodents: house mouse, Norway
rat and black rat. Their relative occurrences, as given by the results of
large-scale trappings, are: house mouse — 90.0%; Norway rat — 9.49%;
black rat — 0.5% (Vereshchagin, 1949) (Figure 206).
Continuing development of tea, tung and citrus plantations will
undoubtedly lead to the cutting of the remaining original forests anda
decrease in the area planted to rice. This will produce the most drastic
impoverishment in all the ecological assemblages of mammals, creating a
situation similar to that described for Colchis.
Sparse juniper—pistachio forests of eastern Transcaucasia
It is known from paleontological and archaeological studies (Petrov,
1939; Vereshchagin, 1949c; Gummel and Yatsenko-Khmelevskii, 1941) that
the dry foothills of the southeasternmost Greater Caucasus in Kabristan
and the Gil'gin-Chai basin, on the Nukha, Kartalinia and Тога plateaus,
on the Kirovabad Plain and southeastern slopes of Karabakh and
in the Gori depression were covered in the Holocene by the so-called
"light forests" consisting of groves of Turkish terebinth (Pistacia
mutica) and junipers (Juniperus polycarpos) alternating with
557
sections of beardgrass steppe. By now there remain only small relict
sections of these forests, heavily damaged by cutting, on the barely
accessible southern cliffs and northern slopes of the Tertiary hills in the
Tsivi-Tskaroi, Samukh, Agdash and Geok-Chai districts. Some preserved
and ritual groves and trees protected by the Moslems, as, for example,
in Kabristan, are also relicts of these juniper —pistachio forests
(Vereshchagin, 1949c). Observations in these residual forests show that the
light forest was inhabited by bear, striped hyena, wolf, fox, badger, stone
marten and red deer.
FIGURE 207. Remains of the riverain and juniper forests in the Tur'yan-Chai ravine
Photograph by author, 1949
The first result of uncontrolled cutting of the forests for firewood and
building materials and of accelerated cattle-grazing was the replacement
of the juniper —pistachio by Christ's-thorn thickets; at a later stage upland-
xerophytic formations of thorny astragali predominated. The pre-18th -
century destruction of vast tracts of sparse forest has considerably
553 restricted the distribution of bear, boar and deer. Increased erosion and
desiccation of the slopes (Figure 207) have driven the populations of steppe
vole and Asia Minor hamster to the northern slopes of the hills and to the
plain areas.
Mountain forests of the Greater Caucasus
In the main, the most recent evolutionary stages of the species' ranges
and of ecological assemblages in the mountain-forest belt of the Greater
558
554
Caucasus are related to the development of the hunting, timber and са е -
breeding industries.
During the last fifty years, bison was exterminated in this zone; the
distribution of deer and panther has drastically declined; in many places
the populations of bear, boar, roe deer, chamois and Caucasian goat have
also considerably decreased.
Extermination of the forests of the lower belt and, later, of the fir
forests of the middle and upper belts has been described by Dinnik (1911) for
the northwestern Caucasus. Commercial cutting of fir and beech forests in
the valleys of the Zelenchuk, Teberda and other rivers began in the 1890's.
The greatest deforestation caused by cutting and cattle grazing occurred
on the crests of the ridges descending in a fan-like pattern to the Taman
Peninsula. The formation of forest glades with mesophytic meadows
increased the populationof pine vole and improved the food supply of boar,
roe deer and deer.
Deeper in the canyons the process of deforestation was accompanied
by orchard planting. Long ago the Circassians planted pear, quince and
medlar along the tributaries of the Afips, Psekups and other rivulets;
and after they were driven out in the second half of the last century, the
pear plantings grew wild and expanded over the canyon bottoms penetrating
to and sometimes into the broadleaf forests. These peculiar pear forests,
described by Maleev (1939a), were rich in food, particularly in the fall,
and highly attractive to marten, badger, bear, fat dormouse, forest
dormouse, boar, deer and roe deer.
Distribution of the mouse-like rodents was most strongly affected by
man's intrusion into the canyon bottoms from which the striped field mouse,
harvest mouse and steppe mouse, like the forest-steppe rodents, migrate
to the highest points as the canyon bottoms are settled.
The synanthropic mountain assemblage of house species initially develops
from the native species, particularly the common field mouse and long-
tailed snow vole. The latter species commonly occurs in cellars and house
basements and inhabits seed beds and vegetable garden environs (Zharkov,
1949).
То the east, the effects of anthropogenic deforestation of the slopes оп
the evolution of biocenoses rapidly increase in magnitude due to the
progressively drier climate. Extensive clearing of beech and pine forests
on the slopes of the Cherek, Urukh, Ardon, Fiagdon and Terek ravines
(see Bush, 1931, 1935) resulted in the development on the southern slopes of
a dry forest-steppe with formations of tragacanth Astragalus inhabited
by hare and migratory hamster, and on the northern slopes of mesophytic
meadows inhabited by mole and common and pine voles.
In Transcaucasia, the beech—hornbeam and chestnut forests of the lower
belt on the Black Sea coastal terraces from Batumi to Novorossiisk were
subjected to particularly extensive destruction.
The once extensive areas of thicket were reduced to small sections of
shrubs alternating with bracken. Factors in the deforestation of the
mountain slopes in Abkhazia are extensive cutting, goat grazing and forest
fires, which prevent new forest growth (Zaklinskii, 1931).
559
555
Extermination of forests and development of numerous settlements
proceeded concomitantly with the planting of orchards, mainly apple and
pear, and decorative plants, and the establishment of parks with planes,
eucalyptus, firs, bamboo and other exotic plants.
Under these conditions a patently synanthropic assemblage of mammals
developed which is poor in species. These aspects are due to the
occurrence of Norway rat in wild habitats, black rat and fat dormouse in
fruit plantings, and European wildcat and jackal near the poultry houses
in villages and in fishermen's and porpoise-hunters'! stations.
One can see additional evidence of the biological reactions of rodents
to the newest agricultural methods in the common field and yellow-heeled
mice, which have adapted to feeding on cork oak plantings (Sviridenko, 1940,
1944), and in the house mouse which feeds on and inhabits seed beds
(Shidlovskii, 1947).
The population of mouse-like rodents in the mountains near Sochi has
greatly increased in habitats formed by broken, fallen trees and branches
left in forested areas cleared during the war. This waste material
provides many additional shelters for the common field mouse, harvest
mouse and pine vole. In addition, hares and foxes migrate to the mountains
along the cleared sections. At the same time there is a decrease in the
Caucasian yellow-spotted mouse population. However, with planned
cutting and complete disposal of the fallen branches and twigs, the number
of mouse-like rodents drops abruptly in the first year following the cutting,
but is restored in the second year when the cleared sections become
overgrown with meadow grasses and shrubs. Under these conditions, the
black rat is added to the assemblage mentioned (Sviridenko, 1945).
Extensive unplanned cutting of forests reduces the abundance of arboreal
rodents — forest and fat dormice — which only completely disappear during
planned cutting. These observations confirm to some extent Shidlovskii's
(1948) conclusion that the distribution ranges of the striped field mouse and
harvest mouse expanded in a southerly direction only in the last few
centuries under the influence of man on the coastal landscapes.
Deforestation of the mountain slopes of Abkhazia and Adzharia produced
a localized population increase in long-clawed mole vole, which occurs in
relict forest glades of the forest zone.
On the spurs of the Rachin ridge and on the Surami, Kartalinian and
Kakhetian ridges, particularly inthe vicinity of Staliniri, Tbilisi and
Mtskheti, extensive cutting of beech forests brought about the most recent
extension of the distribution areaof the Asia Minor hamster and shifts in
the ranges of many forest species. The Mukhran valley, which, in the last
century, was inhabited by bear, lynx, European wildcat, common field
mouse, fat dormouse, boar, roe deer, and deer, was cleared of forests
only in the last 50-60 years. By now the sacred ''witness'"' trees are the
only remains of the beech—hornbeam forests which covered the area in
the past (Figure 208). In recent time the valley has been planted to barley
and corn except for the marshlands which are used for haymaking and
pastures. The plowed sections were invaded from the west by the Gori
steppe vole, European hare, steppe mouse and fox. Only the long-tailed
white -toothed shrew remains of the former assemblage of forest mammals.
Deforestation of the mountain slopes and subsequent cattle grazing in the
vicinity of Tiflis and Mtskheti has promoted desiccation of the slopes and
the eventual replacement of meadow formations by steppe.
560
There has been no reforestation of this region. Overgrazing is
exterminating even the secondary shrub groupings which develop on the
northern slopes following cutting of the beech forests. These shrub
groupings consist of representatives of the genera Crataegus,
Mespilus, Rosa, Prunus, Jasminum, (Cerasus,' Cotoneaster,
Spiraea and Coryllus. On the southern slopes even such shrubs as
Juniperus and Paliurus disappear. Christ's-thorn is more viable,
but eventually this species also retreats up the slope, not only because of
physical destruction, but also because of changes in the physico-chemical
properties of the soil which take place during consolidation, desiccation
and heating by the sun (Sosnovskii, 1915). This anthropogenic process of
progressive aridity promoted the expansion of the Asia Minor hamster,
migratory hamster, steppe vole, steppe mouse, hare and fox. The same
process has reduced the distribution areas of the forest carnivores, rodents
and ungulates. The anthropogenic replacement of the forest by meadow -
forest and meadow landscapes took place under more mesophytic
conditions on the plateaus of South Ossetia (N. A. and E. A. Bush, 1936).
556 In this region the forest assemblage of large species (bear, deer, roe deer,
boar) became impoverished, whereas the populations of common and pine
voles increased.
FIGURE 208. Sacred "witness" trees in deforested and plowed Mukhran valley, inhabited by steppe vole
Photograph by author, 1945
Farther east, on the southern slope, the results of the deforestation
created by cutting are observable in the districts of Kvareli, Zakataly,
561
Nukha, Kutkashen and Shemakha, Cattle grazing on the upper margins
of the forest prevents new forest growth, as the soil is compacted and
desiccated (Yaroshenko, 1956). As the forests were gradually cleared,
cattle-drive routes, several tens of meters in width, developed to connect
the lowland with the highland pastures. These bald belts, which, asa
rule, follow the crests of the longitudinal ridges, serve as migration
routes for foxes and hares which inhabit the lowlands and the highlands.
FIGURE 209. Desiccated mountain slopes resulting from forest cutting on the middle Gerdyman-Chai
River, inhabited by stone marten and common field mouse
Photograph by author, 1949
Migrations of these animals are easily observable in the fall and during
snowfalls. In the districts of Kvareli, Lagodekhi and near Vartashen the
extensive deforestation of the slopes over a long period of time has
probably enabled the common vole to migrate from the highlands and to
inhabit the Alazan-Agrichai valley.
The recent extermination of forests has had a particular and radical
effect upon the composition of the mammalian fauna near the eastern end
of the Greater Caucasus, between Shemakha, Lagich and Konakhkent. In
this area the beech—hornbeam and oak forests of the upper belt were
mostly replaced by mesophytic meadows on the northern slopes of the
gorges or by thickets of low shrubs.
In the middle zone, at altitudes from 700 to 1,300m, the desiccation of
the slopes and development of the steppe assemblages result from the
557 cutting of forests and cattle grazing. The effects of these processes were
compounded by the karst topography (Figure 209).
562
Of the large animals, east Caucasian goat survived in the upper reaches
of isolated canyons, and boar and some bears in relict islands of oak
forests. The pine marten, deer and roe deer have been completely driven
out.
Xerophilous mammals migrate to this region from Shemakha on the
piedmont steppe.
Migratory hamster, steppe mouse and hare occur at altitudes near
1,200m in’ the vicinity of the village of Demerchi. The boundary of the
distribution range of the steppe vole is at a lower altitude — 800-900m.
As awhole, the area is characterized by steppe landscapes on the
plateaus and by mammalian assemblages which are peculiarly impoverished
by an undersaturation of species.
In summary it may be noted that the direct influence of man and the
development of anthropo-cultural landscapes in the forest zone of the
Greater Caucasus result everywhere ina general decline in the populations
of the large game species (bear, boar, deer, roe deer, goat, chamois),
in the disappearance of characteristic forest species (pine marten, fat
dormouse, squirrel), and in a highly differential redistribution of the
mouse-like rodents.
LONGITUDINAL VALLEYS OF THE GREATER CAUCASUS
AND INNER DAGESTAN
The so-called longitudinal valleys are located between El'brus
and inner Dagestan in the east, between the Peredovye range. The
valleys form a line of giant funnels, separated by low ridges. This peculiar
xerophytic warm zone, protected in the north by the ranges as by a
screen, is a refuge for a number of xerophilous and thermophilous animals
and plants. Botanists Krasnov (1893-1894) and Kuznetsov (1909) regarded
this zone as equal in importance to inner Dagestan in the development of
special xerophilous flora. Grossgeim (1948) related the upland xerophytic
formation of this zone to the type of tragacanths, which is characteristic
of the dry uplands of southwest Asia. However, the xerophytic upland
formations with feather grass, wormwood and barberry occur in places
west of the El'brus in longitudinal glacial valleys on the Kuban, the
Teberda, and the Malyi and Bolshoi Zelenchuk rivers, and in the upper
Ingur valley in western Transcaucasia. The longitudinal valleys have been
inhabited by man since the Bronze Age; they were the source area of the
so-called Kobanian culture (Uvarova, 1900), dated as the first half of the
first millennium B.C. The population of these valleys was engaged in
cattle breeding and hunting, and later in agriculture. Toward the end
of the Middle Ages the populations of the large hoofed mammals (boar, elk,
deer, roe deer, Caucasian goat, chamois and bison) became much smaller
on the Peredovye ranges, and they were completely driven out of the
longitudinal valleys.
The summer migrations of deer from the southern slope of the range
were under heavy attack in inner Dagestan, in the canyons of Andi- and
Avaz-Koisu, and on Kara-Koisu. The east Caucasian and Bezoar goats were
driven off the major part of the ranges, particularly off the plateaus, and
563
sheep became extinct in these areas. Undoubtedly, the latter two species
558 had already disappeared from the Gunib by the Caucasian war years of
the middle of the last century.
Human activity enhanced the natural xeromorphic character of the
longitudinal valleys and Dagestan, and is responsible in general for the
Recent wide distribution of upland-xeromorphic vegetation on the northern
slope of the range (Tumadzhanov, 1947). In inner Dagestan, where the
forests were never strongly developed, many centuries of continuous
destruction of the pines in the upper reaches of the tributaries of the Sulak
resulted in their replacement by birch groves (Betula pendula,
B. pubescens). This process has to some extent improved the habitats
and feeding conditions of deer, Bezoar and east Caucasian goats. The
ancient inhabitants of the ravines of Kara-Chai, Kabarda, North Ossetia,
inner Dagestan and Svanetia cleared large areas of the ravine floors of
rocks, and planted gardens and fields, arranging the boulders in circular
piles and in rows. Giant stone walls and small fortresses with towers
were often built of granite blocks and slate (Figure 210).
FIGURE 210. Cereal and potato fields near the village of Ushkul' on the upper Ingur. Pine vole and long-
clawed mole vole inhabit the marginal areas
Photograph by author, 1948
The boulder rows mentioned above, when overgrown with shrubs and
grass, resemble the remains of terminal and lateral moraines. Asa rule,
these ridges, like stone fences in villages and residential buildings, are
sites of concentration of steppe mouse, Ciscaucasian mouse and Caucasian
564
559
snow vole. The snow voles use the heaped stones and crevices in fences
and walls for drying and storing their winter food stocks and for building
nests.
In the Ardon ravine near Zaramag, Turov (1926c) observed a stable
population of migratory hamster inhabiting these stone piles. On the
Baksan, Chegem and Cherek, stone fences and abandoned fields are the
favorite habitats of mountain suslik, a pest of cereal and potato crops.
The mountain water vole concentrates on irrigated fields and in gardens
in Kabarda, Ossetia and Dagestan. In these areas the species develops a
strong fossorial tendency much like the mole rats of Central Asia. This
species is a pest which feeds on the roots of young apple, pear and apricot
trees. Furthermore, irrigation water rapidly fills the holes they make and
creates additional damage by eroding large sections of the slopes.
In inner Dagestan the Asia Minor hamster and pine vole became pests of
cereal crops. Some populations of snow vole on the Gunib plateau have
lost the habit of stocking hay for the winter. Instead, they use the hay
stocked by the villagers in niches and caves in limestone rocks.
The synanthropic assemblage of rodents in the isolated mountain villages
of Dagestan develops from the local species: common field mouse and
snow vole.
The potato, barley and wheat fields in a number of ravines of the Grozny
Region attracted the boar to the woodless mountains, and also contributed
to some increase in the abundance of European hare.
In western Transcaucasia, on the middle and upper Ingur, the clearing
of the fir—maple forests on the slopes resulted in the development of
meadow formations with Pontic azalea (Rhododendron flavum).
Cattle grazing and cutting of shrubs for firewood led to the development
of low, weedy motley grass. Xerophilous species of wormwood grew on
the southern slopes. As the deforestation progressed, bear, lynx, European
wildcat, pine marten, chamois and goat disappeared from large areas.
Continued desiccation of the slopes by the cultivation of cereal crops has
driven out the snow and pine voles, long-clawed mole vole, Caucasian birch
mouse and common field mouse. Open areas of new pastures and fields
were often free of rodents, since the typical field pests (common hamster,
common vole) did not occur in the vicinity.
Such isolated mountain valleys are good examples of presently unoccupied
ecological niches formed inthe process of development of the cultural
landscape.
ALPINE MEADOWS AND ROCK AREAS OF GREATER CAUCASUS
Prior to the appearance and development of nomadic cattle raising, the
landscapes and mammalian assemblages of this zone were practically
unaffected by man. In historical time cattle grazing contributed to the
widening of the meadow zone from below due to the lowering of the upper
forest margins. However, the growth of the upper parts of the meadow zone
was limited by the development of turf meadows which invaded the talus in
the mountain passes. Grazing also promoted the expansion of the Caucasian
rhododendron (Rhododendron caucasicum), not grazed by cattle.
565
This, however, prevented the renewal of birch growth. The upper forest
belt was lowered during the last century by 150-200m at some places in
the Caucasus as the result of cattle grazing, and also by shepherds cutting
timber for firewood. Widening of the alpine and subalpine meadow zone
560 brought about an expansion in the range of the common and pine voles and
of mountain populations of hare and fox. Stone-fenced cattle stations,
trampled-down and well-manured, rapidly became overgrown with dock
(Rumex), thistle (Cirsium), and other species. These localities are
refuge sites of common voles which concentrate on the margins of the
trampled-down areas under the protection of multi-layered vegetation.
Prior to the arrival of shepherds in early summer, bears feed on the green
grass in these places, and boars dig in the ground in search of beetle larvae
and worms.
Overgrazing (i.e., overloading the pastures) inhibits the development
of meadow vegetation. Numerous paths forming a network of steps are
made by the cattle on the slopes. The vertical faces of the steps are broken
by many holes dug by common and pine voles and long-clawed mole voles.
Rainwater and melting snow easily enter the holes, eroding the soil.
Fields are laid out in the uppermost parts of the Caucasian meadows,
overgrown, asarule, by Sibbaldia procumbens and cinquefoil
(Potentilla) and often inhabited by pine voles. Eventually this destroys
the meadow growth over great areas. Large sections of grassy slopes are
displaced by cattle grazing and rodent burrowing. The typically rock-strewn
areas thus formed are used by Caucasian goats, mainly as shelters and
resting places. Intensified erosion from heavy rains brings mudflows
charged with stones. In most regions of the Caucasus mudflows are a new
phenomenon in historical time, developing during the last two centuries of
extensive cattle breeding. Thus, at Nukha the first mudflows occurred in
1889, and at Vartashen in 1895 (Pyl'tsov, 1904). Pyl'tsov wrote in 1904
(p. 268) of the Demir-Aparan-Chai basin in the Kutkashen district of
Azerbaidzhan, where particularly powerful mudflow torrents occur:
"... approximately 25 years ago (i.e., in the 1880's — N. V.) there were
not more than 5,000 sheep on the pastures, whereas now there are more
than 10,000." Overloading the mountain meadows with cattle exposed large
areas of the slopes in the natural amphitheater of this ravine, which in the
past was covered by alpine meadows. Asa result, the east Caucasian goat,
chamois and common vole have been almost completely driven out of the
amphitheater.
Nomadic cattle breeding has also affected the wild hoofed mammals:
deer, goat and chamois. The cattle introduced various diseases into the
mountain zone: glanders, foot and mouth disease, anthrax, brucellosis,
mange, demodecosis, favus and helminthiasis. Mortality among wild
ungulates, particularly goats and chamois, from diseases introduced by
cattle has been recorded from the Caucasian (Dinnik, 1914a; Nasimovich,
1949a, 1949b) and Zakataly reservations (Markov and Mlokosevich, 1935;
Vereshchagin, 1938a).
The intrusions of hunters, shepherds with sheep dogs, and flocks of
sheep and goats on the highland meadows during the three summer months
has considerably affected the natural habits and diurnal cycles of deer,
chamois and goat. The deer and chamois developed the summertime habit
566
561
of remaining during the day in the forest zone and upper forest belt, leaving
for the pastures only at night. This is also true of the Caucasian goat in
the west (Nasimovich, 1949a). In the eastern part, on the southern slope,
the east Caucasian goats form two clearly distinguishable populations in
the summer: one inhabits the passes and rock-covered areas of the range
and does not descend to the forests; the other remains in the forest,
leaving for the upper forest belt and mountain pastures only at night
(Vereshchagin, 1938a).
FORESTS ON THE NORTHERN SLOPES OF THE
LESSER CAUCASUS
In historical time landscape changes were probably minimal on the
northern slopes of the Adzhar-Guri ranges, where large tracts of
broadleaf and fir forests were preserved. These forests still supply
appreciable quantities of bear andpine martenpelts. Nevertheless, a rapid
lowering of the upper boundary of the forest is traceable recently throughout
the area. In the Akhaltsykh District the faunal complex of the dry uplands
of southwest Asia has infiltrated the forest zone.
The rapid displacement of bear, deer, roe deer and marten from the
Aten ravine of the Trialet ridge, causedbythe merciless extermination of
the pine and beech forests since World War I, has been described by Belov
(1924).
In northern Armenia, оп the Somkhet, Bezobdal and Pambak ridges,
small forest islands were preserved in some gulleys (Troitskii, 1934). The
ancient age of human settlements in these regions is substantiated by
archaeological and paleontological materials. Desiccation of the slopes
caused by human activity in the upper parts of the forest zone is particularly
noticeable on southern exposures along the upper reaches of the rivers
which originate on the plateau, as, for example, on the Debeda-Chai. The
section of canyon between the Amamly and Kirovakan stations is an example
of a characteristic interpenetration in a mosaic arrangement of the
distribution ranges of xerophilous species of mammals of the upland steppes
and mesophilous species of the forest zone. Human activity always
promotes encroachment of the xerophilous assemblage of small mammals on
the mesophilous assemblage, following expansion of xerophytic formations.
The distribution ranges of the Asia Minor hamster, steppe vole and
Transcaucasian mole vole show a definite tendency toward extension to
the north down the river valley.
In the lower forest belt and partly in the middle belt, the extermination
of the forest resulted in the development of characteristic formations of
drought-resistant shrubs and meadow-steppe. This enabled the xerophilous
species of the east Transcaucasian semidesert (Asia Minor gerbil, steppe
vole and steppe mouse) to migrate up the ravines.
Anthropo-cultural biotopes of the canyon bottoms (settlements, gardens,
orchards, grain fields) are the main junctions of the ranges of small
mammals of the east Transcaucasian steppes and the upland steppes of the
Armenian upland.
567
562
А number of groups can be recognized in the fairly large collection of
species which is increasing in abundance in the cultural landscape of the
canyons. The first group, associated with buildings and gardens, includes
long-tailed white-toothed shrew, house mouse, common field mouse (in
part) and migratory hamster. The second group, associated with plowed
fields and cereal crops, consists of fox, hare, steppe mouse, pine and
common voles, and, at the boundary of the middle and lower belts, steppe
vole. The middle belt is characterized by an assemblage of rodent pests —
pine and common voles — which occur on second-growth meadows used for
haymaking and pasture. In the upper belt, the bushy thickets of oaks and
tall herbage are inhabited by moles and common and pine voles; roe deer
also commonly occur in these areas. Where marshes have developed at
stream sources and hydrophilous vegetation now predominates in upland
valleys which have been deforested, an increase in the water vole
population occurs.
Although the settlements in the area (e.g., the town of Kirovakan) are
very old, the synanthropic assemblage of rodents still consists mostly of
native species. In 1943 these were identified from a total rodent catch as
follows: house mouse — 86%, common field mouse — 10.3%, migratory
hamster — 3.7% (Sosnikhina, 1946). It is of interest to note that in recently
built-up areas the common field mouse occurs only in the winter-spring
season. Hamsters occur in towns in the winter and Summer, but not in
the spring.
Only small tracts of beech—hornbeam forest were preserved farther
east in the broad amphitheater of the Dzegam and Shamkhor river gorges.
The crests of the longitudinal ridges were completely cleared of forests.
The replacement of the forest assemblages of mammals by meadow animals
is particularly pronounced in the zone between 1,000 and 2,500m.
In this area chamois and boar have disappeared, and the abundance of
wild goat, roe deer and bear has decreased.
The. ranges and populations of the common and pine voles have increased
considerably. Those pine voles which inhabit the meadows which have
developed on deforested lands have evolved into a slightly modified
subspecies, Pitymys daghestanicus nasarovi (Shidlovskii, 1938;
Ognev, 1950).
Because of early human activity in the mountains, the primary beech
forests on the eastern slopes of the Karabakh were preserved only in the
Tertera gorge and on the northern and eastern slopes of Mount Kirs.
Extermination of the forests restricted distribution ranges of the mole,
bear, Persian squirrel, fat dormouse, common field mouse, pine vole
and roe deer.
The slopes which were subjected to timbering, plowing and desiccation
became inhabited by steppe vole, migratory hamster and steppe mouse
which migrated from the foothills. In the Tertera gorge the steppe vole
begins to occur at altitudes of 650-700m; the pine vole at this altitude still
clings to the northern shaded rocky areas and oak—hornbeam groves.
The development of the woodless dry mountain landscapes is even more
clearly pronounced in the south of the Karabakh and on the Armenian
plateau proper.
568
563
ARMENIAN UPLAND AND MIDDLE ARAKS VALLEY
Sections of beech—hornbeam forests, subalpine tall herbage and upland
steppes occur on the high plateaus of the Lesser Caucasus. On the southern
slopes of the plateaus, towards the Araks valley, the mesophytic landscapes
are contiguous to arid thin forests and dry sheep's fescue steppe, which
grades into the zone of thorny Astragalus and Acantholimon. The
upland wormwood— saltwort semidesert is developed in the middle Araks
valley.
Man has inhabited the plateau since the Early Paleolithic (Zamyatnin,
1947; Panichkina, 1948). However, during the second half of the
Quarternary the pronounced changes in the landscapes were due mainly to
tectonic and climatic phenomena.
Even in the postglacial and historical epochs the vegetation of the plateau
has changed considerably.
Sections of upland steppe which developed in the postglacial xerothermal
epoch grew smaller because of somewhat higher humidity and the advance of
the forest phase. The landscapes began to be noticeably affected by man in
the Bronze Age when human activity was immediately reflected in the
distribution of forests and abundance of large mammals. Pedological
studies (Klopotovskii, 1948), geobotanical studies (Kuznetsov, 1900;
Troitskii, 1927, 1939) and archaeological studies (Grossgeim, 1948) all
indicate that the pine and oak—beech forests in Dzhavakhetia covered much
larger areas in the earlier stages of the historical epoch than in the present.
However, deforestation of large areas in Dzhavakhetia and Armenia by
lava flows occurred in the Pleistocene (Yaroshenko, 1941).
In the Lower Holocene the plateau was still heavily inhabited by horse,
bear, deer, roe deer, bison, primitive bull and sheep. The woodless
highland areas were Summer pastures for forest ungulates which sought
refuge from the bloodsucking dipterans of the forest zone.
The very end of the second millennium B.C. is the beginning of the
Bronze Age of human culture in Transcaucasia. Bronze Age culture was
characterized by the development of agriculture and, at a later stage, of
cattle breeding (see Piotrovskii, 1947, 1949).
Remains of the ancient irrigation systems have recently been discovered
on the slopes of Alagez and in the Gekham Mountains. It is known from
Urartu cuneiform writings that the tribes of the Armenian plateau possessed
large herds of cattle in the first millennium B.C. Sardur, son of Agrishti,
twice drove herds (totalling 23,194 head of large cattle and 63,420 of small
cattle) from Yeriarkh (a country to the north of Alagez) (Piotrovskii, 1949).
The development of agriculture and cattle husbandry in the Lake Sevan area
were directly responsible for the beginning of the forest extermination in
that area three thousand years ago.
However, the leafy forests on the Armenian plateau were mostly
exterminated in the 19th century (N.B., 1861; Бай, 1947a).
On the southern slopes of the Armenian upland, in the Armenian and
Nakhichevan A.S.S.R., large areas of drought-resistant thin forests were
exterminated in historical time. The thin forests consisted of silver
hawthorn (Crataegus orientalis), willowleaf pear (Pyrus
salicifolia), juniper (Juniperus polycarpos) and almond
(Amygdalus fenzliana). In the present epoch the only remains
569
564
of these forests are sacred ''witness'' trees which we and Burchak-
Abramovich have described elsewhere (1950).
During recent centuries the development of xerophytic vegetation on the
southern slopes of the Zangezur and Daralagez ranges and of Alagez was
promoted by an expanded nomadic cattle husbandry and by overloading the
pastures. Аза result, the grass cover was trampled down and weeds
appeared in the natural grass assemblages.
As the mountain slopes of the Zangezur became desiccated, the lower
boundaries of some rodent ranges (common and snow voles) were raised
by 350-400m. The occurrence of these animals at lower altitudes in the
immediate past is evidenced by isolated colonies which inhabit mesophytic
sections of the Akulis-Chai gorge, rock-covered areas overgrown by
buckthorn (Rhamnus pallasi, В. spathulaefolia) and dog rose
(Rosa canina) at altitudes near 1,200-1,300m. In this area the colonies
exist next to populations of Persian gerbil. The distribution of the gerbils
gradually extends over dry, rocky slopes which have been overgrazed by
sheep and follows the spread of phrygana higher into the mountains. The
present vertical distribution of assemblages of mouse-like forms is shown
in Figure 211.
Similar cases of mutual interpenetration of the distribution ranges of
the highland (upland-steppe and meadow) and semidesert mammals are
known in the upper reaches of the Arpa-Chai and on the Aiodzor ridge. In
these areas, three species of gerbils (Meriones persicus, M.
blackleri and M.vinogradovi) invade the phrygana zone, whereas the
Asia Minor hamsters concentrate on the boundaries and margins of barley
fields and on wastelands in the mountain-steppe zone. In spite of human
activity, the relatively xerophilous upland-steppe complex displays a
tendency toward expansion (Dal', 1949b).
Common field mouse - 8
Common field mouse -3 Steppe mouse -9
Persian gerbil-3
ae Common field mouse -15 : : :
Snow vole -19 z 3 Common field mouse -21 “Si? Minor gerbiled
Tepe OSes S Vinogradov's gerbil-1
Persian gerbil-6 val inaiahineaata
8 Persian gerbil-12 Steppe yole-12
Migratory hamster-3 ту : Е
р ы Mouse-like hamster - 3 така hamster ee
Mouse-like hamster-2
Ea
Mt, Ilyanlu (2000 m)
4
Аи
Mt. Kapudzhikh
(3917 m)
Mt.
Darry-Dag (1911 m)
FIGURE 211. Zonal distribution of rodents in the middle Araks region (on combined longitudinal profile)
based on 100 trap-nights in October 1947
I—rock-covered areas and meadows with weeds (2,200-2,300m); Il—thorny astragali and rock-covered
areas with dog rose and hawthorn (1,600-1,700m); III—rock-covered areas with buckthorn and almond
shrubs (1,200-1,300m); IV—vineyards in river valleys and on hill slopes (1,000-1,100 m)
The process of xerophytization of the middle Araks valley and the
southern slopes of the Armenian upland developed concomitantly with a
570
reverse process — plantings of tracts of walnut, plane, almond, apricot
and apple trees. Examples of such plantings are the nut orchards in the
vicinity of Yerevan and the large groves of centuries-old plane trees in
the valleys of the Nakhichevan-Chai and Bazar-Chai. There is even some
disagreement among botanists as to whether these groves are of
anthropogenic or natural (autochthonous) origin. Fruit-tree oases in the
valleys of the Alindzha-Chai and Akulis-Chai also cover large areas. The
canal on the right bank of the Zanga River, penetrating granitic rock through
a tunnel several hundred meters long, was built in the 8th century B.C. in
the slave-holding state of Urartu. The canal is still in use for irrigation of
large areas of the upland semidesert. Cuneiform writings of the Urartus
tell us that, at the time it was constructed, a number of groves, gardens
and orchards were planted (Piotrovskii, 1949).
(565)
FIGURE 212. Apricot gardens inhabited by Transcaucasian mole vole and common field mouse in
the Alindzha-Chai valley
In general the planting of trees and development of deserts in the valleys
of the Middle East and central Asia during historical time depended less
on climatic fluctuations than on the development of human culture and,
above all, on irrjgational techniques (see also Berg, 1938). The rapid
disappearance of one of the large piedmont oases occurred in recent times
in the Akulis-Chai ravine, near the town of Ordubad, where several tens of
565 hectares of dense fruit-tree plantings were exterminated by drought within
10-15 years of the destruction of the Armenian village of Akulis by the
Turks sin Lo
In historical time, horse, boar, ASiatic wild ass, deer and Persian
gazelle have been exterminated in the middle Araks valley. There was a
571
sharp decline in the abundance of sheep. The populations and ranges of
steppe vole, migratory hamster and mole vole have increased on the
irrigated lands of the Araks valley and its tributaries. The population of
gerbils and jerboas were displaced to the upper terraces. The garden-
oases (Figure 212) in the valleys and orchards, with their stone walls,
became the habitats of the long-tailed white-toothed shrew, hare, forest
dormouse, common and steppe mice, migratory hamster and Transcaucasian
mole vole. The activity of the Transcaucasian mole vole, which inhabited
the artificially irrigated sections of the Vedi-Chai and Arpa-Chai valleys,
has also increased (Dal', 1948a, 1949b). We observed similar
trends in the Alindzha-Chai valley.
Hydrophilous populations of steppe mice developed on the rice fields
of the Araks valley (Pogosyan, 1945). The increase in mouse-like rodents
was followed by an increase in carnivores (fox, tiger polecat, weasel),
although hunting was well developed.
As a whole, anthropogenic landscape changes in this dry valley promoted
an increase in the number of individuals in small-mammal assemblages
which were in the process of adaptation to the peculiar environmental
conditions of the oases.
Buildings built of raw bricks were inhabited by migratory hamsters,
which became house pests. The numerous churches and monasteries
attracted bats, particularly horseshoe bats, pipistrels and noctules.
566 The original synanthropic assemblage of rodents in small settlements
developed from local species: steppe mouse and migratory hamster. The
results of rodent night-trapping in October 1947 in residential buildings
of some villages in the Alindzha-Chai valley are given in Table 114.
TABLE 114. Species and abundance of rodents (number of individuals per 100 traps)
in small settlements in the middle Araks valley
Khanagya
number
of
individuals
Abrakunis
number
of
individuals
Species
To
Mus musculus subsp. ........
М занеси Гиз tatariews.... 5. -
Cricetulus migratorius.....
Apodemus sylvaticus.......
oa) (eae, байке wise ee
At Leninakan and Yerevan, located on the plateau and connected by
railroad with Baku and Tbilisi, the local residents were displaced by recently
introduced cosmopolitan species. The results of rodent trapping carried
out in these cities in the early 1940's are shown in Table 115, based on the
data of Sosnikhina (1944), and Shidlovskii and Sosnikhina (1944).
Such changes in the original synanthropic assemblage on the plateau and
in small settlements are at present related to highway construction and
development of motor transport.
1704 572
567
TABLE 115. Species and abundance (in %) of rodents in cities of the Armenian
Highland
Leninakan
80
15
о
Species Yerevan
MOUS MSCS SHEN Foon neue oo
Cricetulus migratorius.......
RACES МОТИВ ssasonodgoo
50.5
32.0
17.5
Of the measures taken to enrich the mammalian fauna of the plateau, the
following are noteworthy: the highly successful breeding of nutria on the
Kara-Su rivulet (Sev-Dzhur) near Echmiadzin, which began in 1940
(Sarkisov, 1946b), andthe release of 100 raccoon dogs in 1934 near
Voskresenovka and of 15 female and 4 male axis deer in 1954 in the
Khosrova forest near Garnya; the latter two experiments met with little
success. The breeding of deSman was planned for the lakes of the
Akhalkalaki upland.
FOREST AND UPLAND STEPPE ZONES OF TALYSH
The landscape and biocenotic changes which occurred in Talysh in
historical time are very similar to those described for the Lesser Caucasus.
Hunting with firearms caused the disappearance of tiger and deer from
the area in the 1920's and a considerable decrease in the abundance of
panther and bear. Bison and wild goat, which probably inhabited the
highlands, disappeared much earlier, possibly at the beginning of the last
century. The agriculatural activity of man resulted in large changes in
the composition of the assemblages and the distribution of many species of
mammals.
The cutting of broadleaf forests and cattle grazing were particularly
extensive on the northern spurs of this range. These processes led to the
formation of large, dry forest glades, overgrown on the margins by ferns,
bracken and blackberry. In the upper reaches of the Vilyazh-Chai, plowed
fields of barley and wheat occupy the southern slopes of the ravines.
On the eastern slopes, the upper forest boundary was artificially lowered
by 250-300m, even more in some places. Deforestation and desiccation of
the mountain slopes resulted in the displacement of forest animals from
this area and in a large-scale increase in the distribution areas of the
European hare, steppe vole and migratory hamster, which disperse from
the upland xerophytic steppe. The voles migrate over the deforested crests
and slopes of the ranges to the bottom of the gorges at altitudes of
1,200-1,500m, whereas the old, natural boundary of their distribution was
1,700-1,800m. The rapid drying of the slopes enables the voles to withstand
the high humidity in the deep canyons with an annual precipitation of up to
1,500mm. However, in the zone of the ecological optimum of this species,
precipitation does not exceed 500-550mm, and the upper limit on the
plains is 800mm. This example of variability in the ecological stability
of this small rodent is unique on the Caucasian Isthmus. The infiltration
of xerophilous mammals into the humid mountain-forest zone, traced by
us over vast piedmont and mountain areas of the Caucasus, reached its
578
568
maximum development in this area. However, this infiltration is not from
below, as in many other regions, but from above. In the Lerik and
Yardymly districts of Talysh the high reproductive rate of the steppe vole
endangers the cereal crops almost every year. In June large numbers of
boar are attracted to the upper forest margins by the cereal crops.
The range expansion of the xerophilous species in this area was
accompanied by a decrease in the distribution areas of mole, porcupine,
common field mouse, boar and roe deer (in part).
The assemblage of house mammals in the small settlements located in
the mountain forests is extremely poor, consisting only of the dark long-
tailed shrew, house mouse and Norway rat.
The development of cattle husbandry and agriculture in the upland
steppe of Talysh resulted in the complete extermination of the arid
sparse forests. A similar process occurred in the Nakhichevan
District. At present the 2,000-2,500m upland steppe zone of Talysh is
an arid region of unstable agricultural crops; barley, wheat and rye are
grown at higher altitudes. Wild ungulates (Asiatic wild ass, sheep, Bezoar
goat) probably still inhabited the area 150-200 years ago. The development
of agriculture on the plateau during the last centuries promoted a population
increase in mouse-like rodents (steppe and common vole, Transcaucasian
mole vole, Asia Minor hamster) and in the European hare. The extensive
digging in the alfalfa crops by Transcaucasian mole vole is particularly
noticeable in the Zuvanda valley, at altitudes of 1,800-2,000m. In gardens
and alfalfa fields the mole vole forms piles of earth up to 1m in diameter
and 30cm in height, whereas on virgin lands such piles are never over
25-30cm in diameter.
The assemblage of house mammals in the settlements of this upland is
very similar to that described for the middle Araks valley. Trapping in
the mountain villages of Kel'vyaz and Dyman yields mostly local species.
In a June 1945 trapping, the migratory hamster amounted to 80% and house
mouse to 20% of the catch. Migratory hamster even inhabits garrets and
second floors of residential houses in the summer, withstanding very high
temperatures (Vereshchagin, 1949d). The introduced house mouse and
Norway rat are so far common only in the district centers of Yardymly and
Lerik, which are connected with Lenkoran by highways.
PROTECTION AND ENRICHMENT OF FOREST AND HIGHLAND
MAMMALIAN FAUNA
The known record of game animal protection on the Caucasus goes back
to the Early Middle Ages. However, there is no doubt that protective
measures were in use also in the slave-holding states.
Originally, protection consisted of restricting hunting on particularly
valuable hunting grounds, as, for example, the lands of the feudal barons
in Iran, Armenia and Georgia. Such local rules against hunting on the
lands of beks, khans, shahs, princes and tsars also included the protection
of the natural landscape: groves, reed swamps, water springs, etc.
In ancient Armenia, inthe reign of Khosrou (4th-5th centuries A. D.), oak
forests were planted and fenced and later populated by game animals in
the districts of Garni and Рут and in the Metsamor valley. ''Then the tsar
574
569
ordered all kinds of wild animals to be collected and released within the
enclosure, in order to make this the place of the tsar's hunt, feasts and
recreation'' (Buzand, 1953, p.17). In the 19th century, large sections of
mountain forests were selected and leased for the Grand Duke's hunt.
These were the Kuban hunting grounds, nearly 500,000 hectares in area,
in the canyons of the left tributaries of the Kuban; the Caucasian hunting
grounds in the Karayaz tugai forest (Figure 213) on the Kura, east of
Tbilisi, and the Borzhomi animal reserve near Borzhomi (Shil'der, 1892;
Bashkirov, 1940).
On the grounds of the Borzhomi animal reserve, 500 hectares of
mountain slopes were surrounded by a high fence with entrance traps and
no exits for the animals. The Myzym-Chai and Belokany ravines on the
southern slope of the Greater Caucasus near Lagodekhi were leased by
Count Demidov San-Donato in 1903-1917 for hunting mountain ungulates
(Markov and Mlokosevich, 1935; Markov, 1935, 1938). As these areas were
efficiently guarded and rarely hunted, they became, in effect, game
reservations, where a large stock of large mammals was preserved.
In the early 20th century, the idea of maintaining such hunting reserves
evolved into the idea of organizing state game reservations.
After the Soviet revolution the following mountain and mountain-forest
reserves were established in the twenties and thirties: Caucasian,
Borzhomi, Teberda, Zakataly, Lagodekhi, Gumista, Rits, Telavo-Kvareli
and Saraibulakh. Their total area is approximately 550,000 hectares
(Markov, 1935; Makarov, 1940; Gambarashvili, 1937; Knyazev, 1946;
Vereshchagin, 1947d).
FIGURE 213. Results of one day's boar hunt at Karayazy in 1909 — 48 boars killed
Photograph by E. L. Markov
575
In addition а number of temporary game reservations were established
in the thirties in the foothills and in the mountains. In the 1930-40's the
total number of reserves was 17, comprising a total area of 200,000
hectares. The value of these reserves was negligible, since the protection
of animals, which consisted of prohibiting shooting and trapping, did not
extend tothe forests, which were subject to logging and cattle grazing (Map 97).
The establishment of the Caucasian game reservations stopped the
decline in the large mammal population, particularly goats and chamois,
and later contributed to a gradual population increase in these species.
In view of the prospects for great changes in the landscape of the
Caucasus in the next decades from mineral exploitation and agricultural
expansion, conservation measures must be developed by every available
means. Reducing the size of the Caucasian reservations and permitting
logging on their lands is extremely shortsighted and a false economy.
570 At the end of the 19th century some attempts were made to enrich the
mountain-forest mammalian fauna of the Caucasus with foreign species.
Following Shil'der's recommendations in 1888, 54 European fallow deer
(Dama dama L.) were released in the Borzhomi game reservation.
The animals became well acclimatized, and by 1918 there were several
hundreds of them. Herds of 50-80 head wandered through the area, but in
1919-1920 they were all killed by the local population. Also, in 1888,
7 European red deer (Cervus elaphus L.) from Vilna and 58 from
Austria were released in the reservation in order to "improve the antlers"
of local deer. According to the hunting specialist E. L. Markov, the
number of deer in the reservation was nearly 1,000 by 1918; their numbers
later decreasedto less than a hundred head as the result of unlicensed
hunting.
Ten east and west Caucasian goats were also introduced into the
reservation. The crosses between the two species, and between each of
the two with the local wild goats, produced a number of interesting
malformed hybrids. By 1918 the total goat population of the reservation
was presumably 500, which were wiped out during the Civil War. Such
large numbers of animals as these constantly required increasing food
supplies to maintain them in the reservation even in an environment of
mountain relief. At any rate, the experience of the Borzhomi game
reservation is a spectacular example of the great potential in breeding
both local and introduced species of ungulates in the Caucasus.
More or less planned attempts to enrich the mountain-forest fauna of
the Caucasus were made only in Soviet times. Unfortunately, these
operations were concerned only with fur animals.
It should be noted that the methods by which new species were introduced
into the mountain-forest zone of the Caucasus were strictly empirical.
The theory of acclimatization was not at all well developed and only
occasionally were more or less rational propositions advanced and put into
practice.
Formozov (1930) has recommended that the fir forests of the Caucasus
be populated with common squirrel; Markov (1931b, 1935) has suggested
the introduction of axis deer into the Talysh forests, and of rabbit, skunk,
raccoon dog and chinchilla into the mountains and forests of the Caucasus.
Manteifel' (1943) considered it desirable to introduce the following forms
into the Caucasus: goral, markhor, Pamir argali, chinchilla, Himalayan
576
(571)
571
panda and Australian phalanger. Other proposals concerned the introduction
of mink, raccoon and Siberian polecat into Transcaucasia, and the extension
of the ranges of Caucasian goat and chamois to the Lesser Caucasus
(Vereshchagin, 1939b).
Attempts to introduce fur animals into the Caucasian forests have been
discussed by zoologists (Shaposhnikov, 1938; M. Stepanov, 1939; Kurapova
and Stepanov, 1941; Dal', 1941; Vereshchagin, 1941b,1947c; Rukovskii.
1950).
Some of the more successful cases of acclimatization of fur and hoofed
species in the forest and highland tones of the Caucasus are given below.
Experimental acclimatization of squirrel, raccoon dog, axis deer and
bison has been carried out in the western Caucasus.
FIGURE 214. Altai squirrel in the pine forests on the upper Kuban
In 1937, 134 specimens of Altai squirrel (Sciurus vulgaris
altaicus Serebr.) (Figure 214) were released in the Teberda ravine
in the Teberda game reservation. In 1940 the number of squirrels increased
to 2,400, and in 1941 to more than 3,500 (Kurapova and Stepanov, 1941).
In 1944 squirrels inhabited the forests in the basins of the Aksaut, Kuban,
Teberda, Bolshoi Zelenchuk and Malyi Zelenchuk, feeding on the seeds
of firs, spruces, pines and beech nuts. By 1948 the squirrels appeared
in the Caucasian game reservation and on the southern slope of the range
near Sukhumi, probably bypassing the glacier fields in the upper reaches
of the Laba. Squirrel pelts were being procured as early as the fall of
1948 from the forests of the Bzyb and Kodor basins. The advance of the
squirrels to the east in the zone of the fir—maple forests of Svanetia
5
proceeded very rapidly. They appeared in the Rion basin in 1951. In
1954 gardens in the Gagry area were damaged by squirrels (Rukovskii,
1956)*. Undoubtedly, squirrels will inhabit the entire forest zone of
Transcaucasia in the future, and will probably migrate through the Surami
ridge forests to the broadleaf and coniferous forests of the Trialet and
Adzhar-Imeretian ranges.
At present the expediency of introducing squirrels into the Caucasian
forests is in doubt because the animal is a garden pest, particularly
destructive of nut-tree plantations. The organization of commercial
squirrel hunting is urgently required.
The present (1956) and future distribution ranges of the red squirrel are
shown in Map 95.
Experimental acclimatization of raccoon dogs, raccoons, minks and
Canadian skunks was carried out on the Zakataly-Ismailly lowland in eastern
Transcaucasia. Of these only the acclimatization of raccoons was
successful.
From an initial release of 22 specimens in 1941 west of the village of
Ismailly, the raccoons increased in numbers and distribution, in spite of
unlicensed hunting and extermination efforts in the village gardens. The
entire valley of the Agri-Chai was a suitable habitat for this omnivorous
predator (Figures 215, 216). By 1945 there were 180-200 raccoons
(572)
FIGURE 215. Common raccoon from the forests of the Agri-Chai valley
Photograph by author, 1949
* Rukovskii's geographical data are lacking in accuracy: the occurrences of squirrels which he shows in the
woodless areas near Novorossiisk, Gelendzhik and other cities are incorrect.
578
572
distributed over an area of nearly 250 Ки? (Vereshchagin, 1947с,
1953b). Ву 1949 the distribution area was approximately 850-900km’,
and the number of animals was 800-850. Under favorable conditions,
the rate of distribution and formation of the range on the humid
wooded lowland was very high — as much as 15-20km per year.
Raccoons often tend to concentrate in the vicinity of settlements where
they can always find food: fruit, frogs and poultry.
In order to increase the rate of natural distribution of raccoons
throughout the region, 23 raccoons were caught in August-September
1949 in the Ismaily area. Of these, 18 were released in the forests on
the lower Samur and on the Khachmas lowland (Rukovskii, 1947, 1950).
Additional animals, caught in the Kutkashen and Ismailly areas in 1950 and
1951, were released in the gallery forests on the Terek and left tributaries
of the Kuban, in spite of our warning that they might cause damage to
poultry and game (Vereshchagin, 1953b).
Introduction of other carnivores was not successful. The Canadian
skunks and minks died within one year of unknown causes. A small number
of raccoon dogs still inhabit the lowland forests of the Avtaran valley.
Equally unsuccessful were the attempts to acclimatize raccoon dogs in
the mountainforests of the Teberda game reservation in Transcaucasia and
the Gumista reservation in Abkhazia, South Ossetia and Kartalinia (Map 94).
The animals, as a rule, died of unknown causes during the few years
573 following their release; they often migrated over great distances (Stepanov,
1939; Dal', 1941; Vereshchagin, 19474).
FIGURE 216. Footprints of raccoon, black stork and black partridge on river shoal near the village of Nidzhi
Photograph by author, 1949
Bye)
574
575
During the last few decades nutrias were introduced into the alder-
covered swamps and thickets of the Colchis lowland. The first 48 nutrias
were released in 1932 east of Lake Paleostom. In 1936 the species was
observed in the region of Kobuleti, Supsa, Lanchkhuta and the mouth of the
Khupa, 20-25km from the point of release. In the fifth year of breeding the
distribution area covered at least 500km”. However, the areas actually
inhabited by the animals probably comprised no more than 3,000-4,000
hectares (Vereshchagin, 1941b).
The rapid distribution of the species through suitable marshland sections
of the Black Sea coast, particularly in Abkhazia, was aided by the artificial
distribution of acclimatized animals in the region.
In some areas nutria disappeared as the result of poorly organized
hunting and inadequate protection from enemies in the thicket during frost
and drought periods.
The prospects for free breeding of nutrias in the water bodies of Colchis
will become steadily poorer as the alder-covered marshes are drained and
deforested. However, semi-free breeding may continue to develop
successfully in lakes of the Bebesyri and Inkit type (Vereshchagin,
1950a).
The subalpine meadows of central Dagestan were used for an experiment
in breeding Altai marmot (Marmota baibacina Brandt). In 1934,
113 marmots from the Kosh-Agach District of the Oirotian autonomous
region were released on the high plateau west of the village of Gunib. The
animals occupied areas of rocky calcareous sandstone; some dug
noles under the blocks of limestone on the grassy slopes (Figure 217).
An August 1935 census revealed 47 marmots, none of which were new
generation. At that time the total area of distribution was nearly 25km?,
In 1936 only 15 mature and 2 young marmots were counted (Shaposhnikov
and Krushinskaya, 1939; Lavrov, 1946). In 1937 18 holes with 27 animals
were counted, Our observations in August 1952 showed that the marmots
were distributed over the entire Gunib plateau, an area of approximately
45km?, The densest colonies occurred along the faults on the northern
of the plateau at an altitude of 1,800m (Figure 218). Occasional
groups of young marmots occurred even near the village of Gunib
at altitudes of 1,300-1,400m. The natural distribution of the animals
proceeded westward, in spite of unlicensed hunting. The distribution of
the colonies on the Gunib shows that the major part of the low slopes and
ravine bottoms in Dagestan, covered by ephemeral upland-steppe vegetation,
are unsuitable habitats for marmots. The prospects for artificial
distribution of animals are better on high plateaus similar to the Gunib
and Khunzakh with extensive rock-covered areas in the marginal zone.
The distribution and breeding of marmots is evidently feasible over the
entire upland-steppe zone of the inner valleys of the northern slope of the
Greater Caucasus to El'brus. The better areas for new releases are the
limestone talus at the base of the cuestas on the southern cliffs of the
Skalistyi ridge and in the canyons of the Argun, Ardon, Cherek and Baksan,
as well as in the Ingur and Rion river valleys. Marmots can also be
distributed on the ridges of the Armenian Upland, for example, in the
Lake Sevan area. The present and future distribution ranges of marmots
are shown in Map 95,
580
(574)
FIGURE 217. Habitats of Altai marmot оп the northern drop of the Gunib plateau
Photograph by author, 1952
In 1939, 59 specimens of axis deer (Cervus nippon Temm.) were
released in the Teberda ravine for the purpose of developing a stock of
animals to be used for their antlers. By 1942 the number of deer increased
to approximately 150, but during the German occupation of the area it was
halved. In 1954 a small herd of axis deer was released in the Kusary area
of Azerbaidzhan,
In order to restore the population of European bison in the Caucasus, 5
relatively pure crosses of European and American bison from Askaniya-Nova
(Bison bison L.XB.bonasus L.; European bison strain: Яо)
were released in the Caucasian game reservation in 1940. Ш 1944, а herd
of 11 relatively pure crosses lived in the reservation; the number increased
to 21 (6 males and 15 females) in 1951 and in 1956 to 106. The animals are
kept in open-air enclosures and in open areas under the protection of
mounted shepherds. They are driven in summer to pastures in the sub-
alpine and alpine zones. At present there is sufficient evidence to believe
that the crosses of the two bison species will be viable for distribution over
the entire forest zone of the northwestern Caucasus.
These attempts to introduce new species into the Caucasian fauna show
that distribution ranges develop instantaneously (in the geological time-
sense), provided free ecological niches are available. The development of
new biocenotic relationships when biotopes are saturated with the introduced
species must be a special field of study of applied zoology.
Future work on the redistribution and enrichment of the highland
mammalian complexes of the Caucasus must be organized to take into
581
account the natural conditions and main trends of human activity in different
sections of the region.
FIGURE 218, Altai marmots in Dagestan
576 Planning for future enrichment of the mammalian complex of the
mountain-forest zones of the Caucasus should be based on the following
general theoretical considerations:
1. The mammalian complex of the forest zone of the Caucasus was
recently impoverished as regards large species because ofhuman activity. The
species affected were: European bison, elk, and at some places deer and
chamois.
2. The highland assemblage of hoofed animals and rodents of the Greater
Caucasus is poor in species because it was so long isolated, and because
the relief of the alpine zone is highly diversified and snowfalls are heavy,
particularly in the western part of the range.
3. For the rodents and carnivores of the forest zone, the food regime
is more stable in the western part of the Greater Caucasus than in the
eastern part.
4. А number of valuable species of hoofed mammals and carnivores,
characteristic of the Greater Caucasus, do not occur in the lesser Caucasus
area of the Caucasian District.
Taking these specific features into consideration, more attention should
be devoted to the possibility of restoring the populations of European bison
(or the European-American hybrid), elk and red deer in the Caucasus, and
to the introduction of fallow and axis deer. The food resources of the
forest zone are evidently sufficient for one more fur carnivore of the sable
type. Caucasian goats and chamois should be introduced into the Lesser
582
577
Caucasus. Pine marten and chamois should be bred in the Karabakh,
Talysh and other areas.
In general, artificial transgressions over ecological barriers have great
potentialities for increasing the distribution areas of local and introduced
species.
It would be particularly interesting for studies of speciation and
directed variability to attempt an introduction of the east Caucasian goat
in the west, i.e., into the Adzhar-Imeretian Range, and of the west
Caucasian goat into Karabakh or Talysh and El'brus.
Inner Dagestan must play a special role in the breeding and preservation
of highland ungulates. The mountainous rocky relief and thin vegetation
cover in inner Dagestan prevent an extensive development of agriculture,
gardening or cattle breeding. These factors, as well as the geographic
isolation of the area, could well make inner Dagestan a large reservation
for breeding hoofed mammals which do not depend on the forests.
The following species could be introduced for breeding into this area:
Pamir argali (Ovis ammon L.), bharal or blue sheep (О. nahura
Hodgson), tahr (Hemitragus jemlaicus Smith), markhor (Capra
falconeri Wagner), Barbary sheep (Ammotragus lervia Pallas).
The abundance of Bezoar goat (Capra aegagrus), which once
inhabited the entire plateau, must be restored. Beyond question all these
operations must include the planned extermination of wolves.
The dry uplands of the Lesser Caucasus and Talysh are not as potentially
valuable as Dagestan for the breeding of mountain ungulates, since it is
expedient to use the tragacanth—astragali zone for pastureland and for
plantings of almond, pistachio and olive. Nevertheless, the populations
of Persian gazelle must be restored in the middle Araks valley. The
species could occupy the area between Vagarshapat and Ordubad which is
unsuitable for farming.
The semifree breeding of nutria must be further developed in the zone
of the valley forests in Colchis, Zakataly-Ismailly and the Lenkoran
lowlands, as well as on the Kara-Su rivulets in Armenia.
SUMMATION
The study of the direct and indirect effects of man on the mammalian
fauna of the Caucasian Isthmus during the Holocene shows that this epoch
is a distinct qualitative, anthropogenic stage in the evolution of the fauna.
At this stage the distribution ranges of species, ecological assemblages
and faunal complexes evolved under the direct and indirect influence of
chaotic human activity, which often completely overshadowed the natural
processes of evolution of the landscapes and biocenoses,
Some general and specific regularities in the development of the
mammalian fauna of the Caucasus during the Anthropogenic stage may
be summarized as follows:
1. In the historical epoch the mammalian fauna of the Caucasus was
directly affected by man through unplanned hunting, limited only by thé
contemporaneous level of technological development. The principal cause
of the rapid impoverishment of the Caucasus in large mammals lay in the
583
large-scale game hunts practiced by the feudal barons in the mountains
and foothills and by the nomadic tribes on the plains.
The extinction of lion, tiger, river beaver, horse, Asiatic wild ass,
elk, primitive bull and European bison which inhabited the Isthmus was in
some cases greatly accelerated and in others directly caused by man.
Man was also responsible for the decrease in population and range of bear,
striped hyena, panther, boar, roe deer, red deer, saiga and Persian
gazelle.
The composition and abundance of commercially valuable fur species
was much less affected in historical time. During the last centuries and
decades there has been a drastic decline in abundance and distribution of
only two species: corsac fox and Siberian polecat. The data of the state
fur stations for the last 25 years indicate that various natural causes are
responsible for the fluctuations in numbers in the other fox species. These
generally take place over several years.
The planned extermination of wolf, jackal, little suslik, common and
steppe voles and house mouse, which was begun in Soviet time, has not yet
been successful in decreasing the distribution areas of these species.
2. The process of extinction and reduction of distribution areas of game
mammals was accelerated everywhere by the indirect activity of man as
he altered various types of landscape. The universal extermination of the
forests had particularly grave consequences,
3. The most recent evolutionary stages of species ranges and behavior
can be largely traced to the anthropogenetic effect on the higher nervous
system of some Caucasian game animals. These are reflected in choices
of habitats and often in the complete suppression of ancient instincts toward
horizontal and vertical migrations, diurnal activity, etc.
The main reaction of the hoofed animals to the approach of man and his
domestic animals was escape to protected biotopes such as forests,
mountains and deserts. A number of carnivores and ungulates (bear, jackal
and boar), drawn by new feeding grounds and little threatened by human
pursuit, inhabited areas near human settlements in Azerbaidzhan. Man also
promoted the development of local ecological populations of mammals such
as the ''forest'' and ''rock'' populations of the east Caucasian goat on the
southern slope of the eastern Caucasus and the "house" and '' garden"
populations of boars and European hares in the oases of Azerbaidzhan.
578 The formation of new types of biotopes was reflected in some small
mammals, particularly rodents, by changes in ecological features, and
even in the development of new features. This point is also discussed
below (7).
4, Man-made changes in the landscape have also produced changes in
the more or less stable boundaries of the landscape and zoogeographic
zones. Deforestation has resulted in the replacement of mesophilous
ecological assemblages by advancing xerophilous assemblages.
Thus the Ciscaucasian boundaries of the steppe immigrants shifted
southward into the foothills, whereas the boundary of the southwest Asian
upland-desert district moved north into Transcaucasia. This shift of
xerophilous assemblages and complexes of mammals was particularly
advanced in the east, southeast and south of the Caucasus. In these areas
the advance of the xerophilous forms was aided by the natural zonation and
the drier continental climate. The Caucasian mountain-forest and alpine
584
579
district became a mesophytic refuge — a refuge for mesophilous mammals.
Infiltration of the upland-steppe faunal complex into the zone previously
covered by forests resulted in lowering the northern boundary of the
Armenian transitional district down the canyons on the northern slope of
the Lesser Caucasus, made arid by human activity. A similar picture was
even more strongly developed in Talysh.
5. The development of new ecological assemblages and faunal complexes
of mammals on the plowed and irrigated steppes and semideserts of the
Caucasian Isthmus was made possible by the extinction of large game
species of hoofed mammals (tarpan, Asiatic wild ass, saiga, goitered
gazelle) and rodents (suslik, mole rat, jerboa, gerbil), which do not
tolerate plowing and irrigation. Qualitative enrichment of the faunal
complexes proceeded concomitantly with irrigation and development of the
cultural landscapve. The enrichment was due to the migration of
mesophilous insectivores and rodents from the intrazonal habitats: gallery
forests, reed thickets and adjacent sections of the lowland and foothill
forests. Parallel to these developments, there came an increase in the
populations of some widely distributed carnivores: fox, jackal, weasel,
tiger роеса{.
6. Partial restoration and, in some cases, enrichment of the original
assemblages of forest mammals were the results of the replacement of the
piedmont and lowland forests in Ciscaucasia and Transcaucasia by extensive
plantations of fruit and nut trees wherever they were contiguous to the
forests of the lower mountain belt. The seasonal migrations of large game
animals and the increased abundance of arboreal forest rodents and
carnivores which permanently inhabited the area were the main contributing
factors in the restoration and reorganization of the assemblages of forest
mammals.
7. Assemblages of agricultural pests and synanthropic (house)
assemblages developed initially in each zone from the local species.
The cultural landscapes created many possibilities for the development
of new ecological features and mammal assemblages. The best examples
of these processes are the development of hydrophilous populations of
steppe mouse on rice fields and populations of steppe vole on the unirrigated
wheat and barley fields of eastern Transcaucasia.
The accidentally introduced species play only a minor role in the
development of assemblages of agricultural pests. The Norway rat is an
example of this phenomenon: the species became established in the
vegetable gardens of Ciscaucasia and the Black Sea coast.
8. Development of synanthropic assemblages of mammals occurred
many times and independently in different zoogeographic districts and
sections of the Caucasian Isthmus. The development was based on the
adaptation of native species to various local construction techniques,
according to the evolution of morphological features, ecological possibilities
and needs of individual species. The synanthropic assemblages developed,
and still continue to develop, from the representatives of three orders:
Rodentia, Chiroptera and шзесНуога. Rodents are the group which comes
most into contact with man.
Adaptation of the Caucasian rodents (steppe and mountain-steppe races
of house mouse, common field mouse, migratory hamster, snow vole) to
life in man-made structures is undoubtedly as old as the adaptation of the
cosmopolitan races of house mouse and rat.
585.
Improvement in residential quarters and increase in human population
under the conditions provided by cultural landscapes universally resulted in
the replacement of the native synanthropic assemblage by the cosmopolitan
assemblage.
Introduction of species (Norway rat and synanthropic races of house
mouse) into the cultural landscape and human dwellings proceeded at first
along the Black and Caspianseacoasts, where the animals were introduced
from boats. The development of the house rodent assemblages in the
coastal settlements has essentially been completed. At present the inner
regions of the Caucasus are rapidly being inhabited by introduced
synanthropic rodents. This process is due to the development of rail
and motor transport. The displacement of native rodents from the house
assemblage can best be observed on the high plateaus, for example in
Armenia.
9. The planned reconstruction of Soviet agriculture includes planting
of forests and construction of irrigational networks, organization of forestry
and hunting, and development of settlements. All these measures will
promote rapid evolution of ecological assemblages and faunal complexes of
the Caucasian mammals.
10. In addition to the planned alteration of the landscape, measures
aimed at enrichment of the landscape zones and sections described with
Caucasian and alien game animals will also affect the future development
of the mammalian fauna of the Caucasus.
Particularly interesting and economically important results may be
obtained from the introduction of central Asian highland hoofed species
into Dagestan, and of goats of the Greater Caucasus into the Lesser
Caucasus section of the Caucasian District.
586
580
CONCLUSION
This concludes the review of the main stages of the history of the
mammalian fauna of the Caucasus as known from faunological materials
now available. These stages, developing as marks in the Tertiary,
become progressively clearer throughout the Anthropogene towards recent
times. A generalized summary is given below.
The history of the mammalian fauna of the Caucasus begins in the
Oligocene. The Caucasian mountain-forest complex probably began to
develop in the Middle Miocene. The Hipparion faunal complex migrated
to the plain and foothills of the Caucasus from the south in the Upper
Miocene, when the fauna of the Black Sea, Caucasus and Caspian coasts
did not differ essentially from the west Mediterranean fauna. These
similarities in the development of the mammalian fauna were also
pronounced in the Middle Miocene under increasingly differentiated
conditions of climate and landscape in the Caucasus. Mastodons and
Hipparion became extinct; horses, bulls and elephants appeared.
The transition to the Pleistocene on the Caucasus was also similar to
the transition in Europe. The distribution ranges of large animals, the
index species of the Lower Pleistocene (Elephas trogontherii, horses,
Rhinoceros mercki, giant deer and European bison), covered the
Caucasian Isthmus, i.e., the ranges extended from the Russian Plain to
southwest Asia. The specificity of the faunal development somewhat
increased following the new stages of landscape differentiation and the
increased influence of the Greater Caucasus as an ecological mountain
barrier. Ciscaucasia was infiltrated by the steppe species of the Russian
Plain and semidesert species ef northern central Asia, whereas
Transcaucasia was penetrated by the upland-steppe species of southwest
Asia.
The process of evolution of the mammalian fauna of the Isthmus in the
Quarternary mainly consisted of extinctions and immigrations, rather
than endemic speciation.
The known data indicate that evolution of the species inthe Quaternary —
evolution of mammoths from Elephas trogontherii, evolution of
Caucasian bison from the primitive bison, etc. — did not result in the
qualitative enrichment of the fauna.
The rates of evolution and speciation and the adaptational trends of
Caucasian mammals vary greatly within orders. This indicates that there
are no unifying laws or general regularities in these processes.
The glacial epoch did not produce noticeably northern effects in the
composition of the mammalian complexes in the Caucasus. Undoubtedly,
the glaciations did result, however, in the shifting of distribution ranges
and in increased rates of evolution of many species.
587
581
A number of xerophilous and thermophilous mammals migrated to the
Caucasus from the south in Lower Holocene time because of the development
of xerothermal conditions. Direct and indirect human influence on the
development of ecological assemblages and complexes of mammals in the
plains and foothills increased toward the Upper Holocene, at which time the
fauna became strongly impoverished in large carnivores and hoofed
mammals. However, consideration of all the facts on the extinction of
individual species, changes in their distribution ranges and evolution of
their complexes in the Anthropogene of the Caucasus (Chapters II, III, VI),
attests to the importance of the role played by natural factors in the
above processes. In particular, this applies to those mammals which
originated in the steppes of central Asia.
The newest stage of development of the Soviet economy opened great
perspectives for the restoration and enrichment of the mammalian fauna
of the Caucasus by exterminating harmful species and introducing useful
ones. Further development of the economy and natural resources of the
Caucasus will require the Government to undertake large-scale protection
of sample natural faunal complexes.
Tasks for immediate investigation
The theory and organization of investigations in the near future of the
mammalian fauna of the Caucasus and its history present many far-reaching
and diverse problems.
The collection of bone material at the known and newly-discovered
deposits of Tertiary and Quarternary mammals must, of course, continue.
Systematic observations within the next few years on the eroded Tertiary
and Quarternary beds of the shores of the Tsimlyanskaya, Novotroitskii,
Tshchikskii and Mingechaur reservoirs may reveal bone materials of
great value in future studies.
More thorough investigation must be carried out at such "Ккеу" deposits
as the Tertiary beds near Belomechetskaya, at El'dar and Udabno, in the
Kosyakin quarry near Stavropol and the sand quarries on the northern Azov
coast, in the Lower Quarternary deposits on the Taman Peninsula and in
the Middle and Upper Quarternary asphalt deposits on the Apsheron.
Surveying Pyatigor'e, Karabakh and central Transcaucasia for Paleolithic
sites may produce valuable paleontological material and improve our
present knowledge of the development of the fauna of these regions in the
Anthropogene.
The storage and maintenance of paleontological collections in the
republican, regional and district museums must be improved. Systematic
work on the collections from the regions mentioned would improve our
knowledge of the composition of the ancient mammalian complexes and their
correlation with the known Cenozoic faunas of the western Mediterranean,
Russian Plain and central Asia. Additional identifications of species in the
fossil faunal complexes, supported by paleobotanical studies, will help in
the reconstruction of complete paleographic pictures and development of the
stratigraphy of the Cenozoic continental sediments.
588
Further study of the distribution ranges of Caucasian mammals must
include ecological analyses of their manifestations of endemism, relict
distribution and discontinuities. These analyses are relevant to determinations
on climatic and sea level fluctuations, orogenesis and glaciations.
They must be supported by paleontological evidence. Thus, for example,
our knowledge of the time of infiltration of the central Asian semidesert and
desert species into eastern Ciscaucasia can be improved only through deep
drilling in the Terek-Kuma semidesert and by a paleontological survey of
the northern foothills of Dagestan.
Our study of the specific composition of the Caucasian fauna in the
Cenozoic undertaken to discover general developmental regulations,
particularly in the Anthropogene, left many unsolved problems, especially
with regard to geographic and intraspecific variation.
In this category belong the genetic and ecological interrelationships of a
number of forms described by taxonomists: mole, water shrew and white-
toothed shrew among the insectivores; weasel and bear among the
carnivores; common field and house mice among the rodents; roe deer
among the hoofed mammals, etc.
It appears that these problems cannot be ЕЯ by a single morphometric
method.
No matter how thorough are such works or how well supported by
accepted field observations, many questions remain unanswered: the extent
of interbreeding, pattern of inheritance of characteristics, etc.
The complex relief, variability of the climates and sharply differentiated
landscapes over short distances produce a very complex picture of
geographic variation. In small mammals, rodents being an example, the
geographic variation is tied to the variation in weather conditions in
different years and seasons.
Studies with material collected in different years and seasons which
does not take into account the altitude may only obscure the nature of the
geographic variation.
Thus the investigator — morphologist or faunologist — who has spent a
great amount of labor on measurements can only make more or less reliable
guesses as to the nature and rates of evolution of the forms. We believe
that at this stage of science the faunologist must combine morphological
analysis not only with the usual ecological observations in the field, but also
with experiments under natural or nearly natural conditions.
Organization of such studies is laborious, but nonetheless necessary,
since they will contribute to the solution of the basic problems in the study
of the evolution of organisms.
Further studies comparing the data on the Recent evolution of ecological
and morpho-physiological characteristics of species with paleontological
material will establish accurately the rates of evolution of mammals at
different stages of the Quarternary, and, employing the fossil mammals
for the stratigraphic subdivision of the Quarternary, permit us to foresee
faunal changes in the future.
589
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600
Burchak-Abramovich, М.Т. 1952а. Nakhodka iskopaemogo buivola
(Bubalus sp.) па Kavkaze (Finds of Fossil Bubalus sp. on the
Caucasus).—Izv. AN AzerbSSR, No. 2:63-75.
Burchak-Abramovich, N.I. 1952b. Nizhnepliotsenovyi byk
Urmiabos azerbaidzanicus Bur. iz Youzhnogo Azerbaid-
zhana (Lower Pliocene Urmiabos azerbaidzanicus Bur.
from Southern Azerbaidzhan).— Izv. AN AzerbSSR, No. 6:15-29.
Burchak-Abramovich, М. ТГ. 1952c. Nakhodki overnskogo mastodonta
Mastodon arvernensis Groiz. et Job. v Azerbaidzhane i
vostochnoi Gruzii (Finds of Mastodon arvernensis Groiz.
et Job. in Azerbaidzhan and East Georgia). —Izv. AN AzerbSSR.,
о). 923550}
Burchak-Abramovich, М.Г. 1952а. Pervobytnyi Бук (Bos
mastan-zadei mihi зр. nov.) у chetvertichnoi faune sel.
Binagady na Apsheronskom Poluostrove. Binagadinskoe mestona-
khozhdenie chetvertichnoi fauny i flory, Il. (Bos mastan-zadei
mihi sp. nov. in the Quaternary Fauna of Binagady on the Apsheron
Peninsula. Binagady Site of Quaternary Fauna and Flora, II).—
Tr. Estestv. -Ist. Май. AN AzerbSSR, Vol. 5:181-203.
Burchak-Abramovich, М.Г. 1953a. Materialy К izucheniyu dlinnykh
trubchatykh kostei (diafizov) molodykh osobei binagadinskikh
krupnykh mlekopitayushchikh. Binagadinskoe mestonakhozhdenie
chetvertichnoi fauny i flory, III (Data on Diaphyses of Young
Specimens of Large Binagady Mammals. Binagady Site of Quaternary
Fauna and Flora, III).— Tr. Estestv. -Ist. Muz. AN AzerbSSR,
Мотор
Burchak-Abramovich, М.Т. 19535. Sledy povrezhdenii khishchnymi
zveryamna kostyakh melkopitayushchikh binagadinskogo iskopaemogo
kladbishcha Binagadinskoe mestonakhozhdenie chetvertichnoi fauny i
flory, Ш. (Carnivore Impressions on Bones of Binagady Fossil
Deposits. Binagady Site of Quaternary Fauna and Flora, ПТ). — Ibid.,
Vol 6:278=301.
Burchak-Abramovich, М.Г. 1953c. Materialy К izucheniyu boleznen-
nykh i travmaticheskikh obrazovanii na kostyakh binagadinskikh
iskopaemykh ptits Binagadinskoe mestonakhozhdenie chetvertichnoi
fauny i flory, Ш. (Data on Morbic and Traumatic Formations on Bones
of Binagady Fossil Birds. Binagady Site of Quaternary Fauna and
IMliovez,, Те 5 ВОТ. 6: 310-328
Burchak-Abramovich, М.Г. 1953а. Nakhodki elasmoteriya na
Apsheronskom poluostrove i Yuzhnom Urale (Finds of Elasmo-
therium onthe Apsheron Peninsula and in the Southern Urals). —
Izv. AN AzerbSSR, No. 6:75-90.
Burchak-Abramovich, N.I. 1954. Iskopaemyi nosorog Rhinoce-
ros cf. etruscus Falc.) у doline р. Alazani (Fossil Rhino -
ceros cf. etruscus Falc. inthe Alazan River Valley) — Ibid.,
No. 4:33-48.
Burchak-Abramovich, М.Т. 1957. Iskopaemye byki Starogo Sveta,
1 (Fossil Bos of the Old World, 1).— Tr. Estestv. -Ist. Muz. AN
AzerbSSR, Vol. 11:1-263+XI Tables.
Burchak-Abramovich, N.I. andR.D. Dzhafarov. 1945. Nakhodka
gigantskogo olenya v kirovykh otlozheniyakh Apsheronskogo
poluostrova (Finds of Giant Deer in Apsheron Peninsula Bitumen
Deposits). —Izv. АМ AzerbSSR, No. 10:71-78.
601
Burchak-Abramovich, М. 1. апа В.О. Dzhafarov. 1946. Sus
apsheronicus sp. п. У sostave binagadinskoi fauny (Sus
apsheronicus зр.п. inthe Binagady Faunal Complex).— Ibid.,
2(6):26-57.
Burchak-Abramovich, N.I. andR.D. Dzhafarov. 1948. Ostatki
dikogo kabana iz binagadinskikh kirovykh otlozhenii (Apsheron)
(Wild Boar Fossils from the Binagady Bitumen Deposits (Apsheron)).
— Tr. Estestv. -Ist. Май. AN AzerbSSR, Vol. 1-2:26-37.
Burchak-Abramovich, N.I. andR.D. Dzhafarov. 1949. Leopard
(Pardus pardus tullianus Valenc.) na Apsheronskom
poluostrove (Panther (Pardus pardus tullianus Valenc. )
on the Apsheron Peninsula).— Ibid., Vol. 3:86-104.
Burchak-Abramovich, М.Т. andR.D. Dzhafarov. 1950. Materialy
k izucheniyu verkhnetretichnykh Proboscidea (khobotnykh)
Zakavkaz'ya: rod Mastodon et Dinotherium (Data on Upper
Tertiary Proboscidea of Transcaucasia: Mastodon and
Dinotherium Genera).—Izv. AN AzerbSSR, No. 2:38-54.
Burchak-Abramovich, М.Т. andR.D. Dzhafarov. 1953.
Materialy k kolichestvennomu uchetu kopytnykh Binagadinskogo
mestonakhozhdeniya chetvertichnoi fauny i flory, III (Quantitative
Estimates of Ungulata. Binagady Site of Quaternary Fauna and Flora,
IlI).— Tr. Estestv.-Ist. Muz. AN AzerbSSR, Vol. 6:157-209.
Burchak-Abramovich, N.I. andR.D. Dzhafarov. 1955.
Binagadinskoe mestonakhozhdenie verkhnechetvertichnoi fauny i
flory na Apsheronskom poluostrove. Binagadinskoe mestona-
khozhdenie chetvertichnoi fauny i flory, IV (Binagady Site of Upper
Quaternary Fauna and Floraonthe Apsheron Peninsula. Binagady Site
of Quaternary Fauna and Flora. IV).—Ibid., Vol. 10:89-145.
Burchak-Abramovich, М.О. and Z.S. Ekvtimishvili. 1953.
Nakhodka iskopaemogo nosoroga (Rhinoceros cf.etruscus
Falc. ) iz Kakhetii (selo Tsinandali) (Report on a Fossil Rhinoceros
(Rhinocerus cf.etruscus Falc.) from Kakhetia (Tsinandali
Village)).—Tr. Inst. Zool. AN GruzSSR, Vol. 11:229-236.
Burchak-Abramovich, N.I. and E.G. Gabashvili. 1945.
Vysshaya chelovekoobraznaya obez'yana iz verkhnetretichnykh
otlozhenii vostochnoi Gruzii (Higher Anthropoid Ape from Upper
Tertiary Deposits of East Georgia). — Soobshch. AN GruzsSSR,
6(6):451-457.
Burchak-Abramovich, N.I. andE.G. Gabashvili. 1950.
Nakhodka iskopaemoi vysshei chelovekoobraznoi obez'yany v
predelakh Gruzii (Finds of Fossil Anthropoid Ape in Georgia). —
Priroga, No.9: 10" 712.
Buturlin, S.A. 1934. Losi (Elks). pp. 2-67. — Moskva- Leningrad,
Promyslovye Zhivotnye SSSR, KOIZ.
Byalynitskii-Birulya, А.А. 1917. Scorpiony, 1. (Scorpions, 1).—
Fauna Rossii i sopredel'nykh stran. Paukoobraznye (Arachnoidea),
No. 1:1-224.
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Nakhichevanskoi ASSR (Herpetologic Fauna of Armenian S.S.R.
602
and Nakhichevan A.S.S.R.).— Trudy Biologicheskogo Instituta
Armyanskogo Filiala AN SSSR, Vol.3, Zoological Collection 1:77-194.
Chernyavskaya, S.I. 1956. Sezonnoe razmeshchenie i kochevki dikikh
kopytnykh i medvedya v raione Kavkazskogo zapovednika v svyazi
s raspredeleniem urozhaya i plodov fruktarnikov i orekhonosov
(Seasonal Distribution and Migrations of Wild Ungulates and Bears
in the Region of the Caucasian Game Reserve as Related to the
Yield of Fruits and Nuts).— Byull. Mosk. Obshch. Ispyt. Prirody,
otdel biologicheskii 61(4):7-21.
Cherskii, I. 1891. Opisanie kollektsii posletretichnykh mlekopitayu-
shchikh zhivotnykh, sobrannykh novosibirskoyu ekspeditsieyu 1885-
1886 (Description of Post-Tertiary Mammal Collection made by
the Novosibirsk Expedition in 1885-1886).— Supplement to the 65th
Volume of ''Zapiski Akademii Nauk'', No. 1:1-706. SPb.
Chirkova, А.Е. 1952. Metodika i nekotorye rezul'taty uchetov chislen-
nosti lisitsy i korsaka (Investigations into Corsac and Fox Population
Development and Census). — Metody ucheta chislennosti i geo-
graficheskogo raspredeleniya nazemnykh pozvonochnykh, Izd. AN
SSSR, рр. 179-203. Moskva.
Chishwivyv is ву: 11, 1. №. ам А РЕевакаайе. 1938. Materialiyt k
poznaniyu vrednykh dlya sel'skogo khozyaistva gryzunov,
rasprostranennykh v raione Kaspiya (Investigations of Rodents
Deleterious to Agriculture of the Caspian Region). — Vestnik
Gruzinskogo Gosudarstvennogo Sel'skokhozyaistvennogo Instituta,
1(5):72-93 (In Georgian).
Dal', S.K. 1940a. Nasekomoyadnye i rukokrylye Armyanskoi SSR i
Nakhichevanskoi ASSR (Insectivora and Chiroptera of the Armenian
S.S.R. and Nakhichevan A.S.S.R.).— Zool. Sb. Arm. Fil. AN SSSR,
Мо ев.
, 5.К. 1940. К issledovaniyu vymershikh i sovremennykh zhivotnykh
Sarai-Bulagskogo khrebta (Investigation of Extinct and Recent
Animals of the Saraibulakh Ridge).—Ibid., Vol. 2:27-35.
Dal', Б.К. 1941. Dannye о rezul'tatakh akklimatizatsii enotovidnykh sobak
v Armyanskoi SSR (Data on Acclimatization of Raccoon Dogs in the
Armenian S.S.R.).—Izv. Arm. Fil. AN SSSR, 1(6):121-127.
Dal', 5.К. 1944a. Melkie pushnye zveri Pambakskogo khrebta (Small
Fur-Bearing Animals of the Pambak Ridge).— Zool. Sb. AN
ArmsSSR, Vol. 3:47-69.
Dal', S.K. 1944b. Pozvonochnye zhivotnye Sarai Bulagskogo khrebta
(Vertebrates of the Saraibulakh Ridge).— Ibid., Vol. 3:5-46.
Dal', 5.К. 1945. О nakhozdenii podkovonosa Mekheli (Rhinolophus
mehelyi Matschie) v Armyanskoi SSR (Finds of Rhinolophus
mehelyi Matschie in the Armenian S.S.R.).— DAN ArmSSR,
3(4):97-98.
Dal',S.K. 1946. Novye dannye о zemleroikakh-belozubkakh Armyanskoi
SSR (New Data on White-Toothed Shrews of the Armenian S.S.R.).—
DAN ArmSSR, 4(5):143-147.
Dal', S.K. 1947a. Novye biogeograficheskie dannye ob istoricheskikh
granitsakh lesov v Armyanskoi SSR (New Biogeographical Data on
the Historic Boundaries of Forests in the Armenian S.S.R.).—
Ibid. , 6(3):83-87.
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,
S.K. 1947b. Novyi podvid nochnitsy Natterera iz Vaika (Daralagez)
(New Subspecies of Natterer's Noctuidae from Vaik (Daralagez)).—
DAN ArmSSR, 7(4):173-178.
S.K. 1948a. Pozvonochnye zhivotnye Pambakskogo khrebta
(Vertebrates of the Pambak Ridge). — Zool. Sb. AN ArmSSR,
Vols 5:5-68.
S.K. 1948b. Materialy po vertikal'nomu rasprostraneniyu presmy-
kayushchikhsya, ptits i mlekopitayushchikh v dolinakh rek Zangi i
Miskhany (Data on the Altitudinal Distribution of Reptiles, Birds
and Mammals in the Zanga and Miskhana River Valleys).— Zool.
Sb. AN ArmSSR, Vol. 5:69-86.
S.K. 1949a. Suslik verkhnetretichnykh otlozhenii yuga Zakavkaz'ya
(Suslik in Upper Tertiary Deposits of Southern Transcaucasia). —
Dan ArmSSR,, 11(2):67-71.
S. K. 1949b. Ocherk pozvonochnykh zhivotnykh Aiotsdzorskogo
khrebta (Essay on Aiotsdzor Ridge Vertebrates).— Zool. Sb. AN
ArmSSRae Vols 65-9.
S. К. 1950a. Karlikovyi tur donnykh otlozhenii ozera Sevan (Dwarf
Tur in Bottom Deposits of Lake Sevan). DAN ArmSSR, 11(4):133-
138.
S.K. 1950b. Pozvonochnye zhivotnye pribrezhnoi polosy ozera
Sevan i izmenenie ikh gruppirovok v svyazi so spuskom vodoema
(Vertebrates of the Lake Sevan Littoral: The Ways in which Draining
the Lake Affected Their Composition). — Zool. Sb. АМ ArmSSR,
Violet: 55534
S.K. 195la. K sistematike volkov Zakavkaz'ya (Taxonomy of
Transcaucasian Wolves).— DAN ArmSSR, 14(3):87-92.
S.K. 1951b. Dannye po biologii, rasprostraneniyu i kolichestvenno-
mu sootnosheniyu v stadakh bezoarovykh koz na Urtsskom khrebte
(Data on Biology, Distribution and Population Ratio in Wild Goat
_ Herds on the Saraibulakh Ridge).— Izv. АМ ArmSSR, biologicheskie
2
2
го
>
i sel'skokhozyaistvennye nauki, 4(1):33-40.
S.K. 1952. Rezul'taty izucheniya mlekopitayushchikh iz raskopok
urartskogo goroda Taishebaini (Results of Investigation of Mammals
from the Excavations of Taishebaini, Mt. Ararat).—Izv. AN
ArmSSR, obshchestvennye nauki, No. 1:75-86.
S.K. 1954a. Zhivotnyi mir Armyanskoi SSR (Fauna of the Armenian
S.S.R:.), 1-415 рр. АМ ArmSSR.
S.K. 1954b. Dinamika vidovogo sostava gruppirovok mlekopitayu-
shchikh Tersko-Kumskikh peskov (Dynamics of the Specific
Composition of Mammal Associations in the Terek-Kuma Sands). —
Materialy po Izucheniyu Stavropol'skogo Kraya, No. 6:193-208.
S.K. 1954c. Paleofauna nazemnykh pozvonochnykh iz peshcher
Urtskogo khrebta (Paleofauna of Terrestrial Vertebrates from the
Saraibulakh Ridge Caves).— Izv. AN ArmSSR, biologicheskie i
sel'skokhozyaistvennye nauki, 7(2):61-71.
S.K. 1957. Zakavkazskaya pishchukha (Transcaucasian Pika). —
Zool. Sb. ArmSSR, Vol. 10:17-26.
S.K. and Kh.A. Zakharyan. 1951. Obzor chislennosti gryzunov,
vreditelei sel'skokhozyaistvennykh kul'tur (Population Survey of
Rodent Agricultural Pest).— Izv. AN ArmSSR, biologicheskie i
sel'skokhozyaistvennye nauki, 4(8):757-765.
604
Dement'ev, С.Р. апа А.К. Rustamov. 1946. К voprosu о granitsakh
rasprostraneniya pustynnoi fauny yuzhnogo Ира у Srednei Azii
(Range Limits of Southern Desert Fauna in Soviet Central Asia). —
Izv. Turkm. Fil. АМ SSSR, No. 3-4:147-151.
Dinnik, N. Ya. 1882. Kavkazskii gornyi kozel (Caucasian Mountain
Goat ).— Trudy Sankt -Peterburgskogo Obshchestva Estestvoispitatelei,
Wolke 130859:
Dinnik, М. 1887. On the Caucasian Mountain Goat (Capra caucasica
Guld.).— Ann. Mag. Nat. Hist., June, pp.450-461.
Dinnik, N, Ya. 1896. Kavkazskaya serna i ee obraz zhizni (Caucasian
Chamois and its Mode of Life). — Priroda i Okhota, No. 2:36-59.
Dinnik, N.Ya. 1897. Medved' i ego obraz zhizni na Kavkaze (The Bear
and Its Mode of Life on the Caucasus). — Mater. К Pozn. Faunyi Flory
Ross. Imp., otdel zoologii, Vol. 3:125-169.
Dinnik, N.Ya. 1901. Mlekopitayushchie gornoi polosy Kubanskoi oblasti
(Mammals of the Kuban Mountain Belt).— Ibid., Vol. 5:1-30.
Dinnik, N.Ya. 1902. Kavkazskii olen' (Cervus elaphus maral
Ogil.) (Caucasian deer (Cervus elaphus maral Ogil.)).—lbid.,
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Dinnik, М. Ya. 1909. Kavkazskie kamennye kozly ili tury (Caucasian
Ibex or Tur).—Ibid., Vol. 9:1-47.
Dinnik, H.Ya. 1911. Obshchie zamechaniya o faune Kavkaza (General
Notes on Caucasian Fauna). — Trudy Stavropol'skogo Obshchestva
dlya Izucheniya Severo-Kavkazskogo Kraya у Estestvenno-Istoriche-
skom, Geograficheskom i Antropologicheskom Otnoshenii,
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Dinnik, М. Уа. 1914а. Zveri Kavkaza (Caucasian Animals).— Zap. Kavk.
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Dinnik, N. Ya. 1916. Byvshee bogatstvo Kavkaza dich'yu i istreblenie
ee v poslednee vremya (The Rich Game Resources of the Caucasus
in Earlier Times and Their Recent Destruction). — Nasha Okhota,
Vol. 19:13-24.
Dobrokhotov, V.I. 1939. К nakhodke kamyshovogo kota (Felis
(Chaus)chaus chaus Schr.) v del'te Volgi (Finds of Jungle
Cat (Felis (Chaus)chaus chaus Schr.) in the Volga Delta). —
Nauchno-Metod. Zap. Komit. po Zapov., No. 3:192-193.
Dolgikh, I. 1905. Mnimyi edinorog rimu ili reem Vostoka, ur i tur
Evropy, Bos primigenius paleontologii (Imaginary Narwhal
Rimu or Reem of Orient, Ur or Tur of Europe, Bos primige-
nius)), 1-85 pp. Riga.
Dombrovskii, B.S. 1913. O nakhodke el'darskoi fauny
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Donaurov, S.S. 1949. Nekotorye dannye po biologii lesnoi kunitsy у
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5-38.
605
Donaurov, S.S., У.К. Popov, апа Z.P. Khonyakina. 1938.
Sonya-polchok v raione Kavkazskogo gosudarstvennogo zapovednika
(Fat Dormouse (Glis glis) inthe Caucasian Game Reserve). —
Ibid. , No. 1:227-279.
Donaurov, 5.5. andV.P. Teplov. 1938. Kaban у Kavkazskom
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191-225.
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Samgorskoi ravniny (na gruzinskom yazyke s russkim rezyume)
606
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612
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Gromova, У.Т. 1948. К istorii fauny mlekopitayushchikh Kavkaza (On
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631
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Shidlovskii, M.V. 1940b. Izuchenie gryzunov v Gruzii (The Study of
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Shidlovskii, M.V. 1940c. Ob osobennostyakh rasprostraneniya
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Shidlovskii, M.V. 1951. Gryzuny Yugo-Osetii (South Ossetian Rodents).
=i 0018—2212
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Shidlovskii, M.V. 1954a. Semeistvo peschanok у rodentofaune Gruzii
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635
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Яр: Kavk.) Muzi.)seriyacA, uNo},4:1 205
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Мол
Smirnov, N. 1917. О vidovoi samostoyatel'nosti zakavkazskogo
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1704 636
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640
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Uvarov, B.P. 1917. K faune pozvonochnykh nizov'ev r. Kumy
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Varshavskii, S.N. andK.T. Krylova. 1939. Ekologicheskie
osobennosti populyatsii malogo suslika (Citellus pygmaeus
Pall.) у raznye periody zhizni (Ecological Features of the Little
Suslik (Citellus pygmaeus Pall.) in Various Periods of Its
Life).— Zool. Zhurn., 18(6):1026-1048.
Vasil'yev, Ya. 1896. Predel'nye Пий rasprostraneniya у Kubansko-
Chernomorskom raione krasnogo zverya i ptitsy (Range Limits of
Animal and Bird Game Distribution in the Kuban-Black Sea Area). —
Picicoda-i: OkhotassNonw2a153)5).
Vereshchagin, N.K. 1936. Opyty akklimatizatsii nutrii (Myocastor
coypus bonariensis Rengger) na yuge SSSR (Experiments in
Acclimatizing Nutria (Myocastor coypus bonariensis
Rengger) in the southern U.S.S.R.).— Tr. Azerb. Fil. AN SSSR,
\9129: 15-67.
Vereshchagin, М.К. 1937. Sovremennoe sostoyanie dzheirana
(Gazella sub gutturosa Guld. ) ху Mil'sko-Karabakhskoi stepi
(The Recent Status of Goitered Gazelle (Gazella sub gutturosa
Guld. ) in the Mil'skaya-Karabakh Steppe).—Izv. Azerb. Fil. AN
SSSR, No. 2:155-174.
Vereshchagin, N.K. 1938a. Dagestanskii Tur (Capra cylindri-
cornis Blyth) v Azerbaidzhane (Capra cylindricornis
Blyth in Azerbaidzhan).— Tr. Zool. Inst. AzerbSSR, 9(45):1-70.
Vereshchagin, N.K. 1938b. Mlekopitayushchie Apsheronskogo
poluostrova (Mammals of the Apsheron Peninsula), 1-34 pp. Baku.
Vereshchagin, N.K. 1939a. Dzheiran v Azerbaidzhane (The Goitered
Gazelle in Azerbaidzhan).— Tr. Zool. Inst. Azerb. Fil. AN SSSR
№910: 1109-1122.
Vereshchagin, М.К. 1939. К voprosu о rekonstruktsii fauny
Azerbaidzhana (A Reconstruction of the Azerbaidzhan Fauna). —
Izv. Azerb. Fil. AN SSSR, No. 1-2:161-164.
Vereshchagin, N.K. 1939c. K voprosu ob ekologicheskikh nishakh
i morfologicheskikh adaptatsiyakh (On the Problem of Ecological
Niches and Morphological Adaptations). — Byull. Mosk. Obshch.
Ispyt. Prirody, 48(1):43-52.
Vereshchagin, М.К. 1940а. Novye nakhodki iskopaemykh i sovremen-
nykh mlekopitayushchikh у Zakavkaz'e za period 1935-1940 gg. (New
641
Finds of Fossil and Recent Mammals in Transcaucasia, 1935-
1940).— Izv. Azerb. Fil. AN SSSR, No. 6:108-115.
Vereshchagin, N.K. 1940b. Perspektivy razvedeniya nutrii v Abkhazii
(Prospects for Breeding Nutria in Abkhazia).— Monadire, 1.
Vereshchagin, М.К. 1941а. Stepnoi kot (Felis ornata Gray) v
Vostochnom Zakavkaz'e (Felis ornata Gray in Eastern Trans-
caucasia). — Sbornik Trudov Zoologicheskogo Muzeya Moskovskogo
Universiteta, Vol. 6:305.
Vereshchagin, N.K. 1941b. Akklimatizatsiya nutrii (Myocastor
coypus Mol.) у zapadnoi Gruzii (Acclimatization of Nutria
(Myocastor covpus Mol.) in Western Georgia).— Tr. Zool.
Inst. Gruz. Fil. AN SSSR, Vol. 4:4-42.
Vereshchagin, N.K. 1942a. Domovye gryzuny v Azerbaidzhane i
metody bor'by s nimi (Domestic Rodents of Azerbaidzhan and
Methods of Controlling Them). 1-37 pp. — Baku, Izd. AN AzerbSSR.
Vereshchagin, N.K. 1942b. Katalog zverei Azerbaidzhana (Catalog
of Azerbaidzhan Animals). 1-95 pp.— Baku, Izd. AN AzerbSSR.
Vereshchagin, N.K. 1942c. Usloviya zhizni vreditelya polei-
obshchestvennoi polevki v Azerbaidzhane (Life Conditions of a Field
Pest, the Social Vole, in Azerbaidzhan).—Izv. Azerb. Fil. AN
SSSR, No. 2:85-89.
Vereshchagin, N.K. 1944. Nakhodka polevoi myshi v Azerbaidzhane
(Finds of Striped Field Mouse in Azerbaidzhan).— Priroda,
No. 3:74-75.
Vereshchagin, N.K. 1945a. Zhivotnyi mir Azerbaidzhana (The Animal
Life of Azerbaidzhan).— In book: Fizicheskaya geografiya AzerbSSR,
pp. 233-269. Baku.
Vereshchagin, N.K. 1945b. Novye faunisticheskie nakhodki v Talyshe
(New Faunistic Finds in Talysh).— Priroda, No. 6:67-68.
Vereshchagin, N.K. 1946a. Gibel' ptits ot nefti v Azerbadizhane
(The Loss of Azerbaidzhan Bird Life in Oil [Pools]).— Zool. Zhurn.,
25(1):69-80.
Vereshchagin, N.K. 1946b. Osnovnye ekologicheskie cherty obshchest-
vennoi polevki v polupustynnoi zone Azerbaidzhanskoi SSR (Main
Ecological Features of the Social Vole in the Semidesert Zone of
the Azerbaidzhan §.S.R.).— Tr. Zool. Inst. АМ AzerbSSR,
Vol. 11:144-182.
Vereshchagin, N.K. 1947a. Novye nakhodki khishchnykh i kopytnykh
v binagadinskom asfal'ts (New Finds of Carnivora and Ungulata
in the Binagady Asphalt).— DAN SSSR, 55(3):247-249.
Vereshchagin, N.K. 1947b. Novaya rasa burogo medvedya iz
binagadinskogo asfal'ta (Ursus arctos binagadensis subsp.
nov.) (New Species of European Bear (Ursus arctos binaga-
densis subsp. nov.) from the Binagady Asphalt).—Ibid., 55(4):
351-359,
Vereshchagin, М.К. 1947с. Amerikanskii enot b lesakh Ismaillinskogo
raiona Azerbaidzhanskoi SSR (American Raccoon in the Forests of
the Ismailly Area of the Azerbaidzhan S.S.R.).— Izv. AN AzerbSSR,
No. 5:68-73.
642
Vereshchagin, М.К. 19474. Okhotnich'i i promyslovye zhivotnye
Kavkaza (Game and Commercial Animals of the Caucasus).
3-144 pp.— Baku, Izd. AN AzerbSSR.
Vereshchagin, N.K. 1948. Los' Alces sp. kak nedavno vymershee
na Kavkaze zhivotnoe (Alces sp. (Elk): An Animal Recently
Extinct on the Caucasus).— DAN AzerbSSR, 4(3):124-126.
Vereshchagin, N.K. 1949a. K istorii i sistematicheskomu polozheniyu
losya na Kavkaze (History and Taxonomy of Elk on the Caucasus). —
Ibid. , 65(3):491-493.
Vereshchagin, N.K. 1949b. Zoogeograficheskie karty (Zoogeographic
Charts).—In book: Atlas Azerbaidzhanskoi SSR. p.44. Baku, Izd.
AN AzerbsSsR.
Vereshchagin, N.K. 1949c. Pleistotsenovye relikty Kabristana i
Apsheronskogo poluostrova (Pleistocene Relics of Kabristan and
the Apsheron Peninsula). — Byull. Mosk. Obshch. Ispyt. Prirody,
otdel biologicheskii, 54(4):3-14.
Vereshchagin, М.К. 19494. О proiskhozhdenii krys года Rattus у
Zakavkaz'e (Origin of Rats of the Genus Rattus in Transcaucasia).
—ieriroda, (Neowin 6125.63.
Vereshchagin, N.K. 1949e. K ekologii i epidemiologicheskomu
znacheniyu gryzunov Lenkoranskoi nizmennosti i gornogo Talysha
(Rodents of the Lenkoran Lowland and Talysh Uplands: Their
Ecology and Epidemiological Importance).— Tr. Zool. Inst. AN
AzerbssR;. Vol. 13:115=129°
Vereshchagin, N.K. 1950a. Bolotnyi bobr (nutriya) po razvedenie i
promysel v vodoemakh Zakavkaz'ya (Breeding Commercial Nutria
(Myocastor coypus Mol.) in Transcaucasian Water Bodies).
1-146 pp.— Baku, Izd. AN AzerbSSR.
Vereshchagin, N.K. 1950b. Zimovki i promysel vodoplavayushchei
ptitsy v Azerbaidzhane (Hibernation and Hunting of Waterfowl in
Azerbaidzhan).— Tr. Inst. Zool. AN AzerbSSR, Vol. 14:133-212.
Vereshchagin, М.К. 1951а. Ostatki sobaki 1 bobrov (Mammalia,
Canis, Castor, Trogontherium) iz nizhnego pleistotsena
zapadnogo Kavkaza (Fossils of Dog and Beaver (Mammalia: Canis,
Castor, Trogontherium) from the Lower Pleistocene of the
Western Caucasus).— DAN SSSR, 80(5):821-824.
Vereshchagin, М.К. 1951b. Khishchnye (Carnivora) iz binagadinskogo
asfal'ta: Binagadinskoe mestonakhozhdenie chetvertichnoi fauny i
flory, I (Carnivora from the Binagady Asphalt: Binagady Site of
Quaternary Flora and Fauna, I).— Tr. Estestv. -Ist. Muz. AN
AzerbsSR, Vol. 4:28-126.
Vereshchagin, М.К. 1951с. Mlekopitayushchie (Mammals). —
Zhivotnyi Mir Azerbaidzhana. pp. 84-251. Baku, Izd. АМ AzerbSSR.
Vereshchagin, М.К. 19514. Usloviya massovoi gibeli nazemnykh
pozvonochnykh i zakhoroneniya ikh ostatkov v Zakavkaz'e
(Circumstances of Mass Mortality and of Fossil Deposition of
Terrestrial Vertebrates in Transcaucasia). — Zool. Zhurn.,
30(6):616-619.
Vereshchagin, N.K. 1952. Ostatki zhivotnykh i rastenii v bituminoz-
nykh otlozheniyakh (Remains of Animals and Plants in Bituminous
Deposits). — Priroda, No. 3:122-123.
643
Vereshchagin, М.К. 1953а. К istorii landshaftov Predkavkaz'ya v
chetvertichnom periode (History of Ciscaucasian Landscapes in the
Quaternary). — Izvestiya Vsesoyuznogo Geograficheskogo Obshchest-
va, No. 85, 2:200-201.
Vereshchagin, N.K. 1953b. Opyty razvedeniya novykh vidov
pushnykh khishchnikov v Azerbaidzhanskoi SSR (Breeding
Experiments with New Fur-Bearing Carnivore Species in Azerbaid-
zhan §.S.R.).— Tr. Inst. Zool. AN AzerbSSR, Vol. 16:150-168.
Vereshchagin, N.K. 1953c. Ostatki argaliobraznogo barana Ovis cf.
ammon Pall. iz bituminoznykh sloev Apsheronskogo Poluostrova:
Binagadinskoe mestonakhozhdenie chetvertichnoi fauny i flory, Ш
(Remains of Argali-like Sheep (Ovis cf. ammon Pall.) from
Bituminous Strata of the Apsheron Peninsula: Binagady Site of
Quaternary Fauna and Flora, ПТ). — Tr. Estestv.-Ist. Muz. AN
AzerbSSR, Vol. 6:302-309.
Vereshchagin, М.К. 1953d. Velikie 'kladbishcha"' zhivotnykh у
dolinakh rek Russkoi ravniny (The Great Animal ''Graveyards" in
the River Valleys of the Russian Plain). — Priroda, No. 12:60-65.
Vereshchagin, N.K. 1954. K istorii fauny pozvonochnykh i razvitiya
landshaftov Stavropol'ya v neogene (A History of the Vertebrates
and of Landscape Development of the Stavropol Area in the Neogene).
— Materialy po Izucheniyu Stavropol'skogo Kraya, No.6:169-175 .
Vereshchagin, М.К. 1955. Kavkazskii los' (А1сез alces cauca-
sicus М. Ver. subsp. nov. ya materialy k istorii losei na Kavkaze
(Caucasian Elk (Alces alces caucasicus N. Ver. subsp.
nov. ): Data on Its History in the Caucasus).— Zool. Zhurn.,
34(2):460-463.
Vereshchagin, N.K. 1956. O prezhnem rasprostranenii nekotorykh
kopytnykh v raione smykaniya evropeisko-kazakhstanskikh i
tsentral'noaziatskikh stepei (Former Distribution of Some Ungulates
at the Junction of the European-Kazakhstan and Central Asian
Steppes).— Zool. Zhurn., 35(10):1541-1553.
Vereshchagin, N.K. 1957a. Ostatki mlekopitayushchikh iz nizhnechet-
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Те Zool. Inst .AWN SSSR ом 958.
Vereshchagin, N.K. 1957b. Pleistotsenovye pozvonochnye iz peshchery
Kudaro I v Yugo-Osetii i ikh znachenie dlya razrabotki istorii fauny
i landshaftov Kavkaza (Pleistocene Vertebrates from Kudaro I Cave
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Vereshchagin, М.К. andI.M. Gromov. 1952. К istorii fauny
pozvonochnykh raiona nizhnego techeniya reki Ural (History of
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Vereshchagin, М.К. andI.M. Gromovy. 1953a. Sbor ostatkov
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Moskva-Leningrad, Izd. AN SSSR.
644
Vereshchagin, М, К. ара Т.М. Gromov. 19535. О proshlom fauny
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Veresmoenaciny N. оао. ©1194 9 Ромедеште ptits) 1
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LIST ОЕ ABBREVIATIONS USED IN THE
BIBLIOGRAPHY
Abbreviation
AN URSR
Azerb. Neft.
Khoz.
Azneft'!
Byull. Kavkaz.
Gos. Zapov.
Byull. Komiss.
po Izuch. Chet-
vertichn. Perioda
Byull. Mosk.
Obshch. Ispyt.
Prirody
Dagnarkomzem
DAN SSSR
DAN ArmSSR
DAN AzerbSSR
Ezhegodn. po Geol.
i Mineral. Rossii
Ezhegodn. Zool.
Muz. AN SSSR
Izd. AN SSSR
Izd. AN UkrSSR
Transliteration
Akademiya Nauk Ukrayins'-
koyi Radyans'koyi
Sotsialistychnoyi Respubliky
Azerbaidzhanskoe Neftyanoe
Khozyaistvo
Translation
Academy of Sciences of
the Ukrainian SSR
[in Ukrainian]
Azerbaidzhan Petroleum
Industry
Gosudarstvennoe Ob''edinenie State Association of the
Azerbaidzhanskoi Neftedoby-
vayushchei Promyshlennosti
Byulleten' Kavkazskogo Gosu-
darstvennogo Zapovednika
Byulleten' Komissii po
Izucheniyu Chetvertichnogo
Perioda
Byulleten' Moskovskogo
Obshchestva Ispytatelei
Prirody
Dagestanskii Narodnyi
Komissariat Zemledeliya
Doklady Akademii Nauk
SSSR
Doklady Akademii Nauk
Armyanskoi SSR
Doklady Akademii Nauk
Azerbaidzhanskoi SSR
Ezhegodnik po Geologii
i Mineralogii Rossii
Ezhegodnik Zoologicheskogo
Muzeya Akademii
Nauk SSSR
Izdatel'stvo Akademii
Nauk SSSR
Izdatel'stvo Akademii Nauk
Ukrainskoi SSR
686
Azerbaidzhan Petroleum
Industry
Bulletin of the Caucasian
State Reservation
Bulletin of the Commission
on the Study of the
Quaternary
Bulletin of the Moscow
Society for Natural
Research
Dagestan People's
Commissariat of
Agriculture
Proceedings of the
Academy of Sciences
of the, U.5.8. В.
Proceedings of the
Academy of Sciences of
the Armenian S.S.R.
Proceedings of the
Academy of Sciences
ofthe AzerbaidzhanS.S.R.
Russian Geological and
Mineralogical Yearbook
Yearbook of the Zoological
Museum of the Academy
of Sciences of the
Виа. Re
Academy of Sciences ofthe
U.S.S.R. Publishing
House
Academy of Sciences of the
Ukr.S.S.R. Publishing
House
]2а. Kavk. Мал.
12а. Mosk.
Obshch. Ispyt.
Prirody
Izv. AN SSSR
Izv. AN ArmSSR
Izv. AN AzerbSSR
Izv. AN Kazakh
SSR
Пу, мое И
AN SSSR
Izv. Azerb. Gos.
Univ.
Izv. Azerb. Univ.
Izv. Geogr.
Obshch.
Izv. Kavk. Muz.
Izv. Kavk. Otdel.
Russk. Geogr.
Obshchestva
Izv. Ross. AN
Izv. Russk.
Geogr. Obshch.
Ae, Abibhelkoatyy Е
AN SSSR
Kabgoizdat
KOIZ
Kratk. Soobshch.
Inst. Ist. Mater.
Ка] Чагу
Izdatel'stvo Kavkazskogo
Muzeya
Izdatel'stvo Moskovskogo
Obshchestva Ispytatelei
Prirody
Izvestiya Akademii Nauk
SSSR
Izvestiya Akademii Nauk
Armyanskoi SSR
Izvestiya Akademii Nauk
Azerbaidzhanskoi SSR
Izvestiya Akademii Nauk
Kazakhskoi SSR
Izvestiya Armyanskogo
Filiala Akademii Nauk
SSSR
Izvestiya Azerbaidzhanskogo
Gosudarstvennogo
Universiteta
Izvestiya Azerbaidzhanskogo
Universiteta
Izvestiya Geograficheskogo
Obshchestva
Izvestiya Kavkazskogo
Muzeya
Izvestiya Kavkazskogo
Otdeleniya Russkogo
Geograficheskogo
Obshchestva
Izvestiya Rossiiskoi
Akademii Nauk
Izvestiya Russkogo
Geograficheskogo
Obshchestva
Izvestiya Turkmenskogo
Filiala Akademii Nauk
SSSR
Kabardinskoe Gosudarsten-
noe Izdatel'stvo
Vsesoyuznoe КоорегаЙупое
Izdatel'stvo
Kratkie Soobshcheniya
Instituta Istorii Material'-
noi Kul'tury
687
Caucasian Museum
Publishing House
Naturalists' Society of
Moscow Publishing
House
Bulletin of the Academy
of Sciences of the
WEES Sekt:
Bulletin of the Academy
of Sciences of the
Armenian 8.S.R.
Bulletin of the Academy
of Sciences of the
Azerbaidzhan §$.S.R.
Bulletin of the Academy
of Sciences of the Kazakh
S.9.R.
Bulletin of the Armenian
Branch of the Academy
of Sciences of the
Я. 5.5.8.
Bulletin of the
Azerbaidzhan State
University
Bulletin of the
Azerbaidzhan University
Bulletin of the
Geographical Society
Bulletin of the Caucasian
Museum
Bulletin of the Caucasian
Branch of the Russian
Geographical Society
Bulletin of the Russian
Academy of Sciences
Bulletin of the Russian
Geographical Society
Bulletin of the Turkmenian
Branch of the Academy
of Sciences of the
WhiSs So 199.
Kabardian State Publishing
House
All-Union Cooperative
Publishing House
Brief Communications
of the Institute of the
History of Material
Culture
Mater. К. Pozn.
Fauny i Flory
Ross. Imp.
Mater. k Pozn.
Fauny i Flory
Materialy k Poznaniyu Fauny Materials for the Study
i Flory Rossiiskoi Imperii of Fauna and Flora of
the Russian Empire
Materials for the Study
of Fauna and Flora of
Materialy k Poznaniyu Fauny
i Flory SSSR
SSSR ide log БВ:
Mater. po. Materialy po Chetvertichnomu Materials on the
Chetvert. Periodu SSSR Quaternary of the
Periodu SSSR USS. №.
MGU Moskovskii Gosudarstvennyi Moscow State University
Universitet
Nauchno-Metodicheskie
Zapiski Komiteta po
Nauchno-Metod.
Zap. Komit. po
Scientific-Methodological
Notes of the Committee
Zapov. Zapovednikam on Nature Reservations
Nauchn. Tr. Nauchnye Trudy Erevanskogo Scientific Transactions,
Erevansk. Gos. Gosudarstvennogo Yerevan State University
Univ. Universiteta
NIIS Nauchno-Issledovatel'skii Scientific Research
Institut Svyazi Institute of
Communications
NKTP SSSR Narodnyi Komissariat People's Commissariat
Tyazheloi Promyshlennosti
SSSR
Sbornik Nauchnykh Trudov,
Erevanskii Meditsinskii
Institut
Sovet Narodnykh Komissarov
of Heavy Industry of
the Wro.o: kh.
Collection of Scientific
Works, Yerevan
Medical Institute
Council of the People's
Commissars
Soobshcheniya Akademii Nauk Communications of the
Gruzinskoi SSR Academy of Sciences of
the Georgian S.S.R.
Communications of the
Georgian Branch of the
Academy of Sciences of
tHe WS JS) 5%
Sb. Nauchn. Tr.,
Erevanskii Med.
Inst.
SNK
Soobshch. AN
Gruz SSR
Soobshch. Gruz.
Fil. AN SSSR
Soobshcheniya Gruzinskogo
Filiala Akademii Nauk SSSR
Sotsekgiz Gosudarstvennoe Izdatel'stvo State Publishing House
Sotsial'no-Ekonomicheskoi of Social Sciences and
Literatury Economics
SPb Sankt-Peterburg ow. Freterspure
er Azerb. al: Trudy Azerbaidzhanskogo Transactions of the
AN SSSR Filiala Akademii Nauk Azerbaidzhan Branch
Tr. Biol! Inst.
Arm. Fil. AN
SSSR
SSSR
Trudy Biologicheskogo
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Filial, Akademiya Nauk
SSSR
688
of the Academy of
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Transactions of the
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Armenian Branch,
Academy of Sciences
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Tr. Erevansk.
Zool. Parka
Tr. Geol. Muz.
AN SSSR
Tr. Inst. Zool.
AN AzerbSSR
Tr. Geol. i
Mineral, Muz.
AN SSSR
Tr. Kavk. Gos.
Zapov.
Те. Komiss .“po
Izuch.
Chertvertich.
Perioda
Tr. Paleontol.
Inst. AN SSSR
Tr. Sov. Sektsii
Mezhdun.
Assots. po
Izuch. Chetver-
tich. Perioda
Evropy
Tr. Zool. Inst.
AN SSSR
Tr. Zool. Inst.
AN Gruz. SSR
Tr. Zool. Sektsii
Gruz. Fil. AN
SSSR
Tr. Zool. Sektsii
Gruz. Otd.
Zakavk. Fil.
AN SSSR
Trudy Erevanskogo
Zoologicheskogo Parka
Trudy Geologicheskogo
Muzeya Akademii Nauk
SSSR
Trudy Instituta Zoologii,
Akademii Nauk
Azerbaidzhanskoi SSR
Trudy Geologicheskogo i
Mineralogicheskogo
Muzeya Akademii Nauk
SSSR
Trudy Kavkazskogo
Gosudarstvennogo
Zapovednika
Trudy Komissii po Izucheniyu
Chertvertichnogo Perioda
Trudy Paleontologicheskogo
Instituta Akademii Nauk
SSSR
Trudy Sovetskoi Sektsii
Mezhdunarodnoi Assotsiatsii
po Izucheniyu Chetvertich-
nogo Perioda Evropy
Trudy Zoologicheskogo
Instituta Akademii Nauk
SSSR
Trudy Zoologicheskogo
Instituta Akademii Nauk
Gruzinskoi SSR
Trudy Zoologicheskoi Sektsii
Gruzinskogo Filiala
Akademii Nauk SSSR
Trudy Zoologicheskoi Sektsii
Gruzinskogo Otdeleniya
Zakavkazskogo Filiala
Akademii Nauk SSSR
689
Transactions of the
Yerevan Zoological Park
Transactions of the
Geological Museum of
the Academy of Sciences
of the U.S.S.R.
Transactions of the
Zoological Institute,
Academy of Sciences
of the Azerbaidzhan
а.
Transactions of the
Geological and
Mineralogical Museum
of the Academy of
Sciences of the U.S.S.R.
Transactions of the
Caucasian State Reserve
Transactions of the
Committee on the Study
of the Quaternary
Transactions of the
Paleontological Institute
of the Academy of
Sciences of the U.S.S.R.
Transactions of the Soviet
Section of the Inter -
national Association for
the Study of the
Quaternary of Europe
Transactions of the
Zoological Institute,
Academy of Sciences of
Woe ©. Sais like
Transactions of the
Zoological Institute,
Academy of Sciences of
the Georgian 8S.S.R.
Transactions of the
Zoological Section,
Georgian Branch of the
Academy of Sciences
Git Wore Wa 5.5. В.
Transactions of the
Zoological Section,
Georgian Division ofthe
Transcaucasian Branch
of the Academy of
Sciences of the U.S.S.R.
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Uch. Zap. Azerb.
Gos. Univ.
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Uch. Zap. Sev.-
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Zap. AN
Zap. Kavk. Muz.
Zap. Kavk. Otd.
Russk. Geogr.
Obshch.
Zool, Sh,
Zool Sb. Arima,
Fil. AN SSSR
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Tsentral'nyi Ispolnitel'nyi
Komitet
Uchenye Zapiski Azerbaid-
zhanskogo Gosudarstven-
nogo Universiteta im.
S.M. Kirova
Uchenye Zapiski, Severo-
Osetinskii Gosudarstvennyi
Pedagogicheskii Institut
im. К.Г. Khetagurova
Uchenye Zapiski Severo-
Kavkazskogo Instituta
Kraevedeniya
Zapiski Akademii Nauk
Zapiski Kavkazskogo
Muzeya
Zapiski Kavkazskogo
Otdeleniya Russkogo
Geograficheskogo
Obshchestva
Zoologicheskii Sbornik
Zoologicheskii Sbornik
Armyanskogo Filiala
Akademii Nauk SSSR
Zoologicheskii Zhurnal
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Central Executive
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Scientific Reports of the
Azerbaidzhan State
University im.
S.M. Kirov
Scientific Reports, North
Ossetian State
Pedagogical Institute
im. K.L. Khetagurov
Scientific Reports of the
North Caucasian Institute
of Regional Studies
Reports of the Academy
of Sciences
Reports of the Caucasian
Museum
Reports of the Caucasian
Branch of the Russian
Geographical Society
Zoological Collection
Zoological Collection of
the Armenian Branch
of the Academy of
Seiences of the U.S.S.R.
Journal of Zoology
629 Supplement. Part 1
WIELD AND SO PREV ОЕ ANIMAL (PE LYS
ON THE CAUCASIAN ISTHMUS FROM
WAS) 0.1955
(Graphs 1-23, pp. 692-703)
(643) Part 2
VAP S ОЕ ANIMAL DISTRIBUTION (ON
TE OA CAS US
(Maps 1-97, pp. 705-801)
691
631
Graph 1
Mole
“° Ciscaucasia
25. at eo oie F330 dor 39
1930 1940 1950
Graph 2 Н
И
[8
Г \
1 \
1 \
Jackal 1 \
LY
Ciscaucasia | \
———-~» Transcaucasia | \
SOCDIGORCH Dagestan - |
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/\
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boy i | \- Г
' № aN \ /
i р | у
Г ` :
1 \ |
1 \ |
Г \ м 1
| \ ‘\ 1
я ae cal
oe Nv м rf \!
Vv
ao тт
1930 1940 1950
692
632
Graph 3 Wolf
© Ciscaucasia
—--——, Transcaucasia
5 27 2931 33 $ 37 9 1 94 44 7 9 Я 35
1930 1940 1950
Graph 4
nae Fox
> Ciscaucasia
= SS 2 Transcaucasia
5 27 45 1 3 3 37 39°41 43 45 47 49 St
1930 1940 1950
693
633 Graph 5
Fox
at eee Lranscaucasia
pe и Azerbaidzhan
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1930 1940 1950
Graph 6
Corsac Fox
“~ Ciscaucasia
85-27 28. Ft. FB» 35 FP DI AME ND 05 OP 59
3 5
1930 1940 1950
694
634
Graph 7
Raccoon Dog
Ciscaucasia
—— —— Transcaucasia
Cm
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tne!
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3 441797
41
39
1950
90
Graph 8
Веаг
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53
1950
47 49° SI
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1940
33 $53731
27 29% 91
1930
55
695
Ciscaucasia
Graph 9
Marten
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1\
\
\
\
635
---- Transcaucasia
Dagestan
i nd
SB 27 arn 3 $5 2B 4 @ 6 wea” 5t 53 55
1930 1940 1950
Polecat
“f Ciscaucasia
Graph 10
3 SS
9 449 99
1950
33 3 37 94
1940
27 ain
1930
25
696
636 Graph 11
Siberian Polecat
f Ciscaucasia
Dagestan
»*@ SHI el et A
a7 oly 2 5 ли чб lst 9 55
1930 1940
1950
Graph 12
Mink
ae Ciscaucasia
5 272 93 3 37 B's 8 4 7 WD 51 SB SS
1930 1940 1950
697
637 Graph 13
ri
/ | Badger
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1980 1940 1950
Graph 14
Otter
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638 Graph 15
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т А zerbaidzhan 4 | i! №
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45 47 49 51
2 27 29 131 33 $ 79 41 43
1940 1950
1930
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ase = a asia
9 Transcaucasi
Georgia
*—'- Azerbaidzhan
А бы Dagestan
Graph 16 se
a
§ ". Lf ee . | Matsa:
ie $ 37 3 41 4 45 147949 ! 155
1940 1950
1930
699
639
Graph 17
European Hare
к Ciscaucasia
Е Transcaucasia
29 7229 Bot. 3s8 GB п-т Gh AD 5
1930 1940 1950
ee European Hare
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im
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Graph 20
Nutria ]
———- Georgia Я
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641
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Fat Dormouse
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Graph 22 2890357
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7 Ciscaucasia
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1950
702
642 Graph 23
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INDEX ОЕ COMMON NAMES OF MAMMALS
Antelope 16, 43-44, 55, 60, 66, 71, 240, 355, 460, 545
Ape 38, 55, 123, 160
Argali — see Sheep
Ass: 133, 158, 160, P64,02113242, 214, 222+024,- 22627 20% S41; TTa
Kulan (Asiatic wild ass; onager) 158, 164, 184, 208, 212-213, 216,
224, 229, 23154236 —-237, 239, 243^267-269, 327-3al-7374, 382—-ape,
457, 473, 495, 497, 499, 510, 514, 520-525, 530, 571, 574, 584-585,
782
Badger 123, 131, 139, 143, 147, 163, 175, 223, 236-237, 247, 399-401,
466, 495, 498,504, 516, 525, 536, 538, 550, 554, 558-559, 698, 735
Bat 101, 389, 464, 490, 496-497, 499, 532, 541, 572
Barbastel 466, 490
Asiatic 502, °716
European 502, 716
Horseshoe bat 466, 487, 495, 498, 514, 572
Blasius' 502, 712
Great 22674230. 9500) "alt
Mehely's 500, 711
Simiatl 502: 112
Southern 500, 711
Long-eared bat 502, 715
Long-winged bat 502, 715
Mouse-eared bat 466
Bechstein's 502, 713
Natterer's 502, 714
Sharp-eared 502, 527, 556, 713
Tricolor 502, 713
Whiskered 502, 714
Noctule (great bat) 490, 494, 496, 514, 572
Commion 502. 5278 717
Giant 466, 502, 717
Leisler's 502, 717
Pipistrel 466, 490, 494, 502, 572, 718
Pocketed bat 457, 496, 504, 721
Vespertilio 457, 514, 532, 556
Bicolor 502, 5274 120
Bobrinskii's 502, 720
Kuht's 498, 502. 1532: 00, 619
Nathusius' 502, 719
Northern 466, 502, 721
Ognev's 504, 721
802
Savi's 502, 720
Serotine 466, 504, 527, 721
Bear 44, 99, 108, 131, 143, 147, 149, 160, 170-171, 174, 176, 180,
190120981 21811220: 1221230123025), 2028, 326,359! 3
381, 398-400, 467, 492, 496, 498, 514, 521, 524, 541, 546, 548,
550.554. 1551—5651, 1563, 1565—5609, tora, 0584, 4589
Cave 87, 100, 109—109. Пит, ПИ, 123, 199 19. 159, 155)
160.116. 21061231, 2387240, ‘242! 1259=260,1266,1 2921398, 518, 1726
Hurepean brown 103; 110, 138, 155, 167, 209—210, 226, 236, 236,
260-262, 382, 398, 466, 504, 514, 727
Beaver 130,155, 601) 170, 208%) 210,204) SAG. Sele 402) 523—524,
547, 551, 584, 746
Bumopean ae lise 2 O90 22023236" 258, 1290 “2953810,
506, 514, 746
Taman 746
Bison 18, 60, 74, 87, 91-92, 95, 97-98, 104, 107-109, 111, 114, 116,
LIG=11®, 151-152, 15. WGO$lG62, 10%, 170, WS, 199. ил,
195—199, 201-202, 206, 209,214, 218—219, 1221—2222, 6229; 231;
236—238, 240, 243, 257, 346, 374, 378-381, 447-450, 452, 457, 473,
2991192. 496) i> 21524. 1546—548),. 559,09 5633) 569) 573, orn, 581
Caucasian 160, 373-377, 513, 514, 587, 796, 800
European 374, 581-582, 584, 587
Long-horned 87, 160, 239-240
Primitive 97-98, 105, 192, 328, 345, 355, 373-377, 587, 796
Taman 71, 796
Bours lee 17092, 98, hos) 111 114-118, 132,144, 197, 199, 160: 102,
171-172, 175-176, 180-181, 183-184, 196, 202, 207, 209-211, 218,
220-22 231. 930-238: 290, 243 253 269,271 13:28, 330330), 7346,
434—435, 473, 433, 496, 498, (512,514, 516, 518, 52450531, 534 539,
541, 547-549, 551, 554, 557-561, 563, 565-566, 568, 571, 574—575,
584
Apsheron 784
Taman 330, 434, 784
В 95 sie 113311173:8183, 190-191: 1 I6—19 TAO 2h e208 264285221,
DOS). BS, 580
Primitive 95, 118, 327, 377-380, 430, 451, 473, 492, 496, 512,
514, 523, 569, 584, 797
Game lbs. 60 66074,91, 125, 54 bby 1962120219221, 231, в,
242, 536, 541, 785
Dromedary 158, 219, 495, 497, 785
Giant 785
Cats (for other Felidae see cheetah, lion, panther, tiger) 172, 175, 186,
2]2, 219.257 1266
African wildcat 139, 223, 266, 467, 498, 504, 738
Caucasian wildcat 487 —488, 497, 514
Cave эй" 119. 1128, 8266,6737
Civet 164
European wildcat 118, 138, 143, 164, 218, 266, 274—275, 382, 401,
504, 524 5484 554, 560,6565;9695,804.0 i
Jungle cat 143, 149, 160, 164, 231, 243, 275—277, 383, 467, 482,
506, 514, 517, 531, 534, 538, 541—542. 1544, (5575741
803
Lynx 138, 143, 155, 171, 175, 236, 266, 274, 359, 382, 401—402,
468, 495, 506, 548, 554, 560, 565, 699, 739
Мапи! 456, 466, 495—496, 504, 517, 742
Chamoist 1 PRE 1.26, 1.7958 12451550 153.5 1822 16R58 1G, £167) $1 9R0e tL oae
199-200, 202, 209, 220, 237-238, 253, 365-360, 381—383, 461,
487—488, 491, 496, 512, 559,563, 565-566,°568, 576—577, 582-583, 792
Caucasian 359-360
Cheetah! 13930 143,09 149, 1160,9 2062 6231 ).°239) 266) s2708 (27 7S275.0 407,
467, 498, .506;.414,.523, 63h, 9742
Corsac.— see Hox
Deer 16, 45; 54-55, 58, 60, 66, 71, 87-89, 92, 103=104, 111, 1145 106;
118, 129-131, 144, 154, 156-157, 160, 166, 171—éa, 93-196; eRe tee.
184, 187, 190-191, 194-200, 202, 204, 206-207, 209-210, 218-220,
223—224. .240—241, 243, 253, 269, 272, 345-346, 380-361, /435-—438,
451-452, 458, 487-488, 492, 495-496, 498, 522, 524, 530-531, 541,
546-548, 554, 557, 559-561, 563-564, 566-567, 569, 571, 571, 573,
56. 8582
Axis 573, 576-577, 581-582, 800
Fallow 159, 164-165, 241, 267, 336, 341-342, 473, 576, 582, 800
Giant. 89, 98, 104,148, 155, 167,206, 238-239, 241-242): 342-344,
518. 587
Red 54, 100, 125, 153. 160, 1622 1732 18472022 207% 0-2:
216. 218, 220, 222, 231, 236-238) 35-341, (365, ЯЗВА
473,a486,° 5402.9 528215512 658. 8769825 Sst A0ks7
Bac. 71, 118-119, 4164. 180—181, 485, 192, 196, 98 -— 2ООзалае.
206-207, 209, 231, 236-237, 239-240, 243, 253, 274, 346-351, 355,
359, 374, 381-383, 473, 486, 488, 490, 493, 496, 512, 514, 522, 524,
530, 546-549, 551, 554, 557, 559-561, 563, 567-569, 574, 584, 589
Maral (Caucasian) 437, 475, 493, 514
Desman 50, 246, 250—251, 514, 550, 573, 706
Common 500
Dog 168, 172-174, 176, 178, 184-186, 205, 210, 212, 219-223, 229,
РАЗ 25. 253, Wis). Dae, 25a), alos
Siberian red 240, 722
iTamian: 1, 028
Jackal 139, 143, 164, 166, 207-209, 223-224, 231, 236, 239, 243,
251—252 ,,255; 281» 374, 383, 468, 482, 492 9495,6400. 004 Sas: SINC,
521, 531, 534, 536, 538, 541-544, 550, 552, 557, 560, 584—585, 688,
aie
Wolf-95, 100—111.-117, 131, 1847136; 1389) 1435 153) 163.
175—176; 210, 223, 236, 238—239, 252-253, 255-256, 218.7 3261836,
350, 359, 366, 374, 380-381, 389-390, 452, 466, 495, 498, 504, 5108
521, 524-525, 530—532, 536, 539, 541-542, 552, 558, 583—584, 693,
из
Dolphin 68, 171, 297, 236, 520
Dormouse
Common 469, 493
Fat 181, 208, 218, 264-265, 469, 486, 490, 506, 546, 548-550,
552,554, 557, 559-560, 563,9568,8700, 147 t
Forest 132, 145, 181, 216, 469, 506, 525, 527, 534, 548-550,
552.554. 556; 5595550219725 6аь
804
Dromedary — see Camel
Elasmothere — see Rhinoceros
Miephant 50, 155. 60 Gon Ob me 1— Oo. 920 95% 980 151-52 1156: "160,
328, 414-417, 419-429, 447, 452, 460, 475, 487, 778-779
Southern 58, 60, 66, 71, 240-241, 415-417
Bist 9, 104, Sil 78,6164, 170, t90—191 194—199, 202" 206" 23H:
236, 238, 243, 382, 440, 451-452, 457, 479, 487, 493, 512, 523-524,
546, 548, 563, 582, 584
Caucasian 175, 237, 343-349, 440, 514, 788
Ermine — see Weasel
Hox 91 В 95, hit wilis Sa, | MSOR 0143 „А, OSS. M163 1 > = 6"
РОО 212, 223, 23298. 281, 253—256, №8]. ЗИ 330, 385386,
390—398. 451-452) 466,991, 495,498, '504,'521` 524, 526/528,
850. (51.1528: 1550, SS 558, 560—562, 5606. 568512
585, 693—694, 724
NOHO BA WAS), We, 105, 29. Bos 208, 392
Corsac 15. L390, 143, 149-1268, 55.60: MIs al 7 OL 239) 8255256,
991—592, 397—399) 958.499) 5081519, 15, 52-525, 527. 5869,
125
Red 253
Silver 798
Gazelle 16, 42-43, 54, 71, 104, 114, 164-165, 272, 277, 473, 571,
583—584
Coitered 54, 1, (96), 133. 1164. 165} 1182} 12110=213. 216: 1218)
229, 289, Aol, Al, 209, 248, 20%, 988. Salo 354, S56, 959, 91, 319,
382-383, 457, 475, 482, 495, 497, 499, 512, 514, 520, 522, 524,
530—532, 538—541, 545, 585, 791
Gerbil 69, 77, 133, 163, 166, 223, 239, 247, 254, 410, 452, 461, 469,
493, 495, 517, 534-536, 544, 572, 585
Arazdayan 495, 510, 768
Asia Minor 302-303, 382, 469, 472, 478, 493, 499, 510, 541,
567, 570, 767
Great 183, 472, 508, 517, 765
Midday 912, 481,508, 1516 511% 15281 570, 765
Persian 160, 239, 248-249, 303, 381-382, 469, 472, 478, 481,
493-495, 508, 535, 766
Red-tailed Libyan 132, 139, 145, 149, 303-305, 381, 410, 469,
473, 498-499, 508, 531, 536, 538, 541-542, 765
Mamearisk 472, 510, 516—511, 570, 766
Vinogradov's 469, 478, 493, 510, 768
Giraffe 36, 39, 42-43, 71
Glutton 231, 238, 262, 400, 731
Goat 16, 91-92, 100, 103, 107-108, 111,114, 119, 124, 149, 155,
192—108, 167, 102=10. 118, 164, 190=ИФ. 190, 1908. 199-202.
204—205, 207—212, 216-217, 220, 222, 224, 229, 272, 359, 367,
BAS). бо. AGil, 407, BSD, 565, 595-500, DO, 990
Bezoar (wild) goat or ibex 36, 47, 116—117, 164, 167, 202
216, 2205 223, 226-225 oo2 5 DD), oO = ОЗ, 981—383, tose
461, 475, 477, 487, 490, 492-493, 495, 497, 512, 521, 524,
563—564, 568, 573, 576, 583, 794
805
East and west Caucasian goats 159-160, 162, 190, 198-200,
202-203, 208-210, 237-238, 360-367, 381, 443-447, 452, 461, 473,
487-491, 493, 512, 559, 563-564, 566-567, 574, 576-577,
582—584, 793
Markhor 473, 576
Hamsters 49.7116, 2400, 905, AT 7 ALO: Уаз 9bS3, P1495 PGs, SieU ks ie
183, 186, 210, 223, 237, 370, 382, 406-407, 451, 468, 486-487,
490, 495, 513-514, 524,°528,°548,°550,) 5681063
Asia Minor (golden) 108, 117-118, 145, 149, 160, 164, 167, 204,
229, 231, 238, 240, 289, 299-302, 381, 457, 469, 477, 493,
495-497, 499, 508, 548, 558, 560-561, 565, 567, 570, 574, 762
Black 526—527, 548
Common 160, 238, 382-383, 470, 479, 499, 508, 514, 524, 527,
548, 763
Migratory (gray) 132, 139, 145, 149, 202, 382, 407, 469, 495,
499, 508, 516, 526-527, 532, 536-537, 548—5495 559, 561,563,
265. 1568.:510,71502 ole, (595, 64
Mouse-like 227, 469, 478, 494-495, 508, 570, 765
Transcaucasian — see Asia Minor
Hare 55, 139, 171=1 72" 75) 186s) 2085211015223, 82265 (256-2 eae
214.1 277, 336, 404, 468, 492, 513, 524, 527, 532, 534,8538ee04de
544, 48h a5 0) 1554 19557 POV 562 № 56655681512
Blue 119, 133, 149, 163, 242, 468
European 145, 231, 279-281, 403, 468, 487, 495, 499, 506, 516,
526, 1536, 7549-550. 6560,!) 1565 21574715848 100 СТА
Tnttle earth 70; 481, 50625261528 1580015730752
Rabbit 460, 468, 542-543, 576, 799
Hedgehog 138, 179, 246, 384, 463, 496, 527, 534
European (common) 172, 216-218, 229-230, 246, 380, 382, 463,
HOOPS 149 b 16.534, 1541: 152240705
Long-eared 149, 163, 186, 223, 230, 239, 247, 382, 387, 463,
495, 497-498, 500, 514, 516, 524, 526-527, 530, 53275961598 1105
Hippopotamus 55, 155, 164-165, 294, 460
Horse) 18, 45,50, 584160, 68" 66: 2745192595, 04, WOM, ОЭ у lien
116911 Sip 125.212%10129-5131, 01338014435 148:5453=154)0158=060,
162, 166-167, 170, 172-174, 176, 178, 184-186, 190-191, 205,
207—208; ;210=2:1 27 214; 92:16)12195224 1231 62884241) 25152072,
323—328,. 345, 431=433, «473, ) 49578497, 0499, °508,° 5245053681569)
57. 584, 597: 195
Southwest Asian 512, 783
Tarpan 325-926, 457, 43; sol 0), 514517. 6523 —525-8 58508785
Нуепа>59, 715. 625; 131, 134, 1537 2421, 2565 2582 2705 496
Cave 119, 139% 143,/°148,) 153,163, 238i 288=239), 924255257,
518, e5%1
Bi'dar 725
Spotted 143, 154
Striped’ 143,164," 210,: 216, 12305 23:7 ,,243,~257.<258/) 270, 09935
486, 492, 497, 499, 504, 514, 531, 558, 584, 725
Ibex — see Goat
Jackal — see Dog
806
Jerboa 17. 132—133, 139, 160, 210, 238, 405—207, 2410, 4521346169,
БН, 1524-9527, 536,544 5721585
Great 145, 147-148, 155, 160, 163, 238, 406, 470, 473, 480, 499,
500. ЭЛЯ, 526-58, Wail
Northern three-toed 470, 481, 506, 514, 528, 753
Small five-toed 145, 160, 163, 456, 470, 481, 495, 499, 506,
5. S26, 528, Dols 150
Thick-tailed three-toed 470, 473, 481, 506, 514, 528, 753
William's (mountain) 145, 149, 469, 493, 495, 497, 499, 506,
Hotkey 5364554. 152
Kulan — see Ass
Lemming 149, 163
Steppe 163, 319-320, 383,510," 514," 526-527, 768
Leopard — see Panther
ом 1143.16 4 LOL 2G 20-22 1,925,243 9206-200, .527,"302, 468,
29211497: C499." 504, 5049523, 531.541, 5645 197
Cave "Поп — see Cat
Macaca 162, 230, 240, 382, 461
Markhor — see Goat
Mammoth 18, 87-89, 91, 96-98, 104, 118, 124, 149, 161, 163-164,
167, 238-239, 266, 414, 416-419, 421, 429, 447, 518, 587, 780
Manul — see Cat
Maral — see Deer
Marmot 18, 123, 155, 175, 240, 282, 469, 487-489, 514, 524, 744
Altai 580-582, 799
Bobak 469
Caucasian 282, 744
Steppe 282
Memon 119.139. 155, 210. 224. 226. 263—265, 916, 400, toad),
567, 696
Pine 138, 143, 162, 164, 263-265, 382, 400-403, 457-458, 467,
ABT. ADs. BOL, 540, 554 508. HOD, SOY, 999, 129
Stome 130113 223, 226, 237, 263-264, 382, 40115601492,
95 49, 504 51 525 558, 562, 1728
Mastodon 39, 43, 49, 63, 69, 414-415, 587
Long-snouted 28, 71
Swamp 26, 71
Ио 139 401, ou, 470. АЭИ 950 5 550, 5е-5Т9, GUN, 953
American 798
Mole 49, 138, 230, 238, 246-250, 310, 317, 381, 383-384, 387-388,
452, 457-458, 490—491, 525, 548, 539, 568, 574, 589, 692," 706
Eastern 247, 500
Caucasian 247-248, 250, 381, 387, 496, 498, 500, 514, 526
Nole wat 55. Lino 193, 470, 514. 524. 526-527, 548.565 19595
Giant Russian 183, 508, 516, 754
Lesser 469, 477, 497, 508, 754
Russian 470, 506, 754
Mouflon — see Sheep
Mouse 133, 202-203, 223, 410, 469, 552
Caucasian yellow-spotted 408-409, 469, 508, 546, 554, 560,
564, 760
807
Birch 91, 488
Caucasian 360, 458, 486, 491, 493, 496-497, 506, 565, 749
Northern 470, 506, -749
Southern 506, 514, 526, 749
Meld 115, 17282168213. 22s oOo ao
Broad-toothed (Asia Minor) 408, 457, 461, 469, 476, 508, 759
Common 145, 408-409, 452, 469, 491-492, 508, 525, 527-528,
534, 541, 546, 548—550, 552, 554, 556—551,. 559-:560,, 5625/4405,
56891510-—572, 2574: 1585: 158951761
Striped 149, 298-299, 382, 470, 479, 499, 508, 514, 525-526,
550, 559-560, 758
Yellow-necked 408, 470, 508, 514, 759
Harvest 149, 457, 470, 508, 514, 559-560, 759
House 44, 172, 223, 309, 383, 407, 469, 487, 497, 508, 527, 532,
537—538, 552,.554,.556—557, 560, 574, 584, 586, 589, 757
Steppe 139, 145, 229, 516, 524, 526-528, 532, 534-536, 544;
548—550, 554, 557, 559-561, 563-564, 567-568, 570, 572, 585
Noctule — see Bat
Nuiria 293, 528; 535, 543-544, 551, 573, 580, 503, 697.9799
Onager — see ‘Ass
Otter 11655639077, 2138, © 143, 0272, +236, 04925 498. 504 4.524 5594;
698, 736
Ох 92. 343
Musk 149, 163, 473
Panther 123, 143, 166, ‘184, 208, 210, 226, 239—240 266.262.
271-274, 278; 359, 401," 403, 457, 488, 504, 514, 524, ese, 541,"
552. OOS, Sta. .534. 739
Р1ка"49. ‘55. „066, 223—224, 231, 464, 497
Afghan 744
Armenian 468, 506, 744
Small 163, 238, 468
Steppe 744
Pipistrel — see Bat
Polecat 170, 161, 504, 524, 59680731
Siberian 18, 149, 163, 2384°480;:504,.514, 524, (526 =5215 545
577. 696, 131
Tiger 143, 149, 164, 223, 239, 382, 400, 457, 4611435, 4985
504, БЫ: 516, 526.536, 558, 548) 550, SE. 95685, 730
Porcupine 55, 116, 118-119, 123-124, 139, 145, 149, 160,162. 166,
231, 237, 239-240, 272, 290, 469, 473, 487, 492, 510, 556—551,
574, 585
Indian crested 145
Rabbit — see Hare
Raccoonol?=579
Common 529, 550. 518. 198
Dog 528—529, 550, 573, 576-573 26954198
Rat 469, 552
Black 162, 209; °211, 216, 231, 236-231, 294-295-297. .581
469, 473, 487, 492, 498, 508, 532, 534, 536-538, 541, 544, 552, 554,
556-=55а 550:'755
808
Norway 145. 202, 208—209, 213; 216 23], 295—298 328 2388341473"
487, 495, 497, 499, 508, 516, 527, 532, 535, 537-538, 548, 551-552,
556-557, 560, 574, 585-586, 756
Water 77, 180-181, 524, 529, 531, 534-535, 702
Reindeer 92, 119, 149, 163-164, 238, 242, 262, 346, 451, 473
Rhinoceros 28, 36, 39-40, 43, 49-50, 55, 58, 60, 74, 125, 127,
131-133, 144, 149, 154, 165, 167, 239-241, 328, 429-430, 781
Elasmothere 781
Woolly rhinoceros 91, 97, 149, 164, 231, 239, 266, 294, 429-430,
781
Sanoawhone 18. 24 171,292, 104 134, 611-151 ole 153, 155.160, 162.
Ио UIST, 231, ABI, 238—239, Oe 936: 31. 346) 352.
354—356, 381-382, 440-443, 457, 475, 480, 499, 512, 514, 517, 522,
524—525, 527, 544-545, 584-585, 790, 800
Seal 33, 40, 44, 48, 60, 68, 116, 123-124, 166, 184, 210, 212-213,
229. 237, 310—373, 520
ЭБЕРТ А, 119) 11555 MiGs. №0; I= aby 108-183-134. 186.
139—190, 196—202, 209—212, 220-224, 226, 229, 281, 352, 310=3173,
536, 564, 569, 574, 583
ие аи 4. 118. 155, 160. 167. 238-240 6310-3503; 499% (566% 525
576, 583, 795
Armenian mouflon 159, 223, 237, 239, 272, 370-373, 461, 475,
477, 489, 495, 497, 512, 795
Shrew 138, 230, 246, 461, 463, 497, 500, 525, 589, 708
Caucasian 388, 496, 509, 514, 708
Common 388, 457, 463, 487, 500
Lesser 457, 463, 486, 491, 498, 500, 514, 526, 707
Water 463
White-toothed 149, 223, 226, 230, 381, 388, 463, 534, 536,
541, 544, 589
Gray i2bo5. S00 a2 er 10
Little 463, 495, 500, 526-5271, 709
Long-tailed 138, 229, 388-389, 491, 495, 498, 500, 530,
535—536, 554, 556, 560, 568, 572, 710
Persian 497, 500, 710
Pygmy (Etruscan) 463, 500, 710
White-bellied 463, 500, 514, 516, 526-527, 709
Skunk 18, 576
Canadian 578-579
Squirrel 55, 282, 468—469, 491, 554, 563, 576-578
Persian (Caucasian) 469, 506, 546, 554, 697, 744
Red 578, 799
Susinik 18, 76, 168, 118. Zils. BAD. 241, 1232 HOS) DAHA, Os.
585, 703
Asia Minor 231, 289, 382, 404-405, 469, 477, 497, 506, 744
European 404
Large-toothed 163, 469
Е Mao. lor tls, 183, 186, 2040 237-233)! 282—259,
381-382, 404-406, 480, 487, 506, 514, 517, 524, 526-521, 549,
584, 745
Swine 50, 59, 172, 174
Giant 71
Warthog 49, 164
809
Tapir’ 49-50, . 71
Таграп — зее Ногзе
Tiger 231, 243, 266, 268, 210-271, 388,.468; 492, 'а95„`497, ‘499,
504, 523, 531, 541, (559 ;578,, 584,.131
Saber-tooth 44
Tur 151; 155,¢167,°192}"196, 218,. 226; 2377 239;> 241," 374,- 381
445, 524
Vespertilio — see Bat
Vole 133, 145, 149, 160, 210, 223, 231, 240, 384, 410, 461, 486,
491, 527, 536, 546, 549—550, 774
Asia Minor snow 242, 316-318, 411, 469, 495-497, 510, 771,
Caucasian snow 162, 180-182, 229, 231, 313-316, 360, 381-383,
411-412, 452, 457, 469, 476, 486-488, 491, 493, 496-497, 514,
565. 51011585782
Common 139, 145, 148, 161,7172, .180, .809-313; 323; 423, 457}
469-470, 487, 493, 497, 510, 524, 526-528, 548, 550, 554, 556,
559, 561—562: 565—566. 568, 150, 574, .584, 715
Common red-backed 319, 382, 470, 479, 487, 493, 495, 510, 776
Long-tailed snow 318-319, 381, 469, 476, 486-487, 490, 510,
5265, 559% 773
Mole. 132, 139, 149, 160,, 163, 167,. 183, 249,. 322. ‘410,, ‘464,
521, eo lee
Northern 480, 495, 510, 514, 548, 769
Pine 139, 116! 1680—1582 2429 305) SPOS sts. S60. sole our
412-413, 457, 469, 476, 484, 489, 491-492, 510, 547-548,
552, 559-661, 564-566, 568, 774
Promethean (long-clawed mole vole) 320-322, 360, 381, 383, 458,
461, 469, 476, 486-491, 510, 560, 564-566, 769
Steppe 76, 145, 149, 186, 210, 216-217, 242, 248, 254, 289,
305-311, 381-383, 414, 472, 492, 495, 497-499, 510, 514, 516,
526, 531, 534-538, 541-542, 544, 549-550, 554, 556-558, 560-561,
563, 567-568, 570, 572-574, 584-585, 776
Transcaucasian 145, 149, 242, 469, 478, 490, 495, 510, 567,
571—572, 574, 769
Water 148, 180, 238, 277, 290, 319, 322-323, 410-411, 452, 469,
490—491; 510, 527, 535, 554, 1565, “S558; 70
Warthog — see Swine
Weasel 138, 143, 223, 316, 401, 488, 504, 516, 526-527, 538, 544,
272. 585: 589, 138
Ermine 504, 731
Whale 22, 40, 48
Toothless 32
Wolf — see Dog
810
INDEX OF LATIN NAMES OF MAMMALS*
Aceratherium 41, 43
— filholi 27*
— incisivum 52
— persia 44
— transcaucasicum 41*
— zernovi 44
Achtiaria 41
— borissiakii 39
— expectans 44
Acinonyx jubatus 141, 145*,
283, 469, 663+
Alactagulus acontion 473, 668+
Alces alces 105, 108*, 109, 117, 118, 440,
475, 686+
— — caucasicus 180, 202, 205, 347. 686+
Alcicephalus neumayri 45
Allactaga 127, 406 ‘
219,
154, 280—
— elater 146*,
530, 667+
— dzhafarovi 141
jaculus 408, 473, 668+
jaculus bogatchevi 141, 146*, 153,
408, 668+
— williamsi 444, 154, 219, 227, 228, 230,
408, 473, 530, 550, 668*
— — dzafarovi 146*, 668+
Ambelodon fricki 32
Amblycastor caucasicus 52, 665+
Amphicyon 31, 32*
Anancus arvernensis 52, 54*, 71, see taxon
Mastodon arvernensis
Anchitherium aurelianense 31
Anthracotherium 27
Antilopinae 45, 47
и 409
— agrarius 149, 301, 473, 544, 671+
flavicollis 149, 4A0*, 473, 672+
fulvipectus 409, 410*, 473, 550, 672+
mystacinus 409, 473, 672+
sylvaticus 144, 149, 153, 182, 183,
185, 193, 206, 208, 218, 219, 295, 228,
230, 4A0*, L73F 530, 544, 547, 550,
566, 673+
— — arianus 534
Archeoceti 26
Archidiskodon —see Elephas
Artiodactyla 27, 39, 41, 43, 44, 47, 48,
52, 57, 59, 60, 61, 63, 69, 71, 74, 96,
123, 127, 141, 193, 233, 333—383, 434,
474, 476, 530, 547, 550, 692+
Arvicola 4141, 677+
230, 408, 473,
a
* [This index has been reproduced photographically from the Russian.
Asterisks denote appearance in figures, crosses in maps
to are at the left-hand side of the text.
(Supplement, Part 2). ]
— terrestris 111%, 141, 149, 453, 183,
185, 186*, 195, 227, 228, 230, 326, 444,
474, 534, 547, 550, 677+
— — cubanensis 412
— kuruschi 412
— — persicus 412
— — tanaiticus 412
Barbastella barbastellus 467, 650+
— darjelingensis 467, 650+
Benaratherium callistrati РИ *
Bison 61, 63, 67*, 71, 77, 100, 404, 118,
448*, "690+
— Бопазиз 122, 152, 241, 475, 692+
— — bonasus 448*, 449*
— — caucasicus 202, 205, 448%,
690+
— longicornis 448*, 449*
— priscus 96, 97*, 99,
115*—117. 449, 124*,
376, 448*, 449*, 690+
— — deminutus 92, 96, 103, 448*
— longicornis 94, 92
— — schoetensacki 63, 77, 93, 100, 376,
448*, 449*
sp. (tamanensis) 448*, 449*, 690+
Bos 59, 64, 91, 100, 118, 434, 161
= mastan-zadei 141, 152%, 454, 380,
691+
— minutus 163, 380
— primigenius 100,
475, 691+
— taurus fossilis 100
trochoceros 162*, 380
Во 41, 71, 354—383
Bubalis 69
449*, 376,
ТОО ча, 312,
127, 161, 163,
152, 160, 161, 380,
Calomyscus bailwardi 230, 473, 675*
Camelopardalis 39
— attica 45
— parva 42
Camelus 59, 134
— dromedarius 163, 223*,
— knoblochi 161, 685+
— kujalnikensis 60, 63
Canidae 52, 54*, 255— 260
Canis 60, 99, 141, 144, 160, 654+
— aureus 144, 155, 170, 210, 212, 227, 255—
256, 469, 530, 547, 550, 653+
— lupus 60, 96, 97*, 105, 108*, 109, 115*,
117, 118, 119, 122, 427, 144, 153, aie
685+
The Russian page numbers referred
811
179, 208,
550, 654+
— — apsheronicus 141, 145*,
— tamanensis 63, 67*, 654+
Сарга 105, 127
—- > ees 225, 227, 370, 475, 689+
caucasica 99, 109, 141%, 142, 115*,
1147—118— 119, 122, 124*, 127, 193,
194*, 205, 209*, 211, 212, 444*, 415,
689+
— caucasica 446*
— severtzovi 446*
cylindricornis 202, 205, 212, 443, 444,
446*, 475, 689+
dinniki 444
— severtzovi 444
Capreolus 104
— capreolus 108*, 109, 117, 118, 122, 127,
130*, 177, 183, 190, 193, 202, 205, 208—
209*— 210, 217, 351, 475, 547, 550,
687+
— — pygargus 687+
Саргоушае 119
Carnivora 31, 39, 41, 44, 52, 57, 59, 60,
63, 74, 123, 127, 144, 233, 255—283,
391, 468, 476, 530, 534, 547, 550, 691+
Castor fiber 117, 211, 293, 473, 665+
-- tamanensis 63, 67*, 665+
Castoridae 47, 293—297
Cervidae 41, 47, 55*, 63, 71, 101, 338—354
Cervus 57, 59, 60, 61, 66, 71, 91, 161,
686+
elaphus 60, 93, 96, 97*, 99, 108*, 109,
iit, 115*—118, 122, 424%, 127, 134,
152 1153.62. 175, 177, 181, 183, 190,
193 194%, 195, 208, 210, "217, 222’
225, 227, 338, 437*—439*, 475, 550,
686+
— binagadensis 141
— maral 161, 202, 205, 438
nippon 575, 692+
pliotarandoides 685+
Cetacea 31.39.43
Cetotherium 37, 39, 43
— meyeri 28
— priscum 43
Chalicotheriidae 34
Chalicotherium pentelici 44
Chionomys — see Microtus
Chiroptera 123, 127, 233, 390, 466, 476,
533, 534
Citellus citellus 292, 473, 665+
— pygmaeus 149, 191, 193,
473, 665+
— — boemii 406
— — musicus 406
— — planicola 406
— — satunini 406
Clethrionomys glareolus 323, 474, 680+
Cricetus 53*, 408, 674+
— cricetus 77, 99, 105—108*—109, 111*,
182, 183, 185, 186%, 190, 406, 473,
674+
Cricetulus migratorius 154, 182, 183, 185,
206, 218, 219, 225, 227, 228, 200) 231:
473° 5307533, (534,566, 674+
227, 256—258, 469, 530, 547,
654+
194*, 285,
812
— — argiropuloi 141,
— Crocidura 52
—lasia 466, 674+
leucodon’ 143, 228, 389, 466, 530, 533,
534, 550, 647+
russula 141, 143, 154, 218, 219, 225,
2271,228} 230, 466, 550, 647+
— caspica 390
gildenstaedti 389, 390, 530, 532—
533—534, 544
— monacha 390, 547
suaveolens 466, 647+
zarudnyi 466, 647+
Crocuta 129, 655+
— eldarica '39, 41*, 655+
— spelaea 96, 97*, 99, 127, 141, 144, 145*,
153, 160, 261, 655*
Cuon 127, 653+
Dama 685+
— dama 692+
— mesopotamica 162*, 170*, 345
ВИ» 26, 39, 210,
elphis 69, 175
Desmana 52, 53*, 250,
— moschata 254—255,
Dicerorhinae 32
Dicerorhinus 43, see taxon
— caucasicus 31, 683+
— etruscus—see Rhinoceros etruscus
— orientalis 52, 56*, 683+
Dicrocerus elegans 31
— salomeae 4
Dinocyon thenardi 52, 54*
Dinotherium 41,
giganteum 39, 42, 48
Dipus sagitta 473, 669+
Dyromys nitedula 141, 149,-185, 218,
219, 228, 473, 530, 544, 547, 550, 666+
Elasmotherium 60
— caucasicum 63, 77, 431
— sibiricum 100
Elephas 74, 91, 681+
— antiquus 59, 63, 68, 101, 415, 682*
—- armeniacus 74, 164
lyrodon 59
mammontoides 681+
meridionalis 59—63, 67*—71, 74, 78,
93, 94, 161, 415, 423*, 681+
primigenius 91, 92, 94, 96119727799;
401, 103, 104, 128, 162, 418, 421*,
682+
trogontherii 63, 69, 77, 78, 91—94,102,
164*, 674+
645+
466, 645+
Rhinoceros
103, 160, 161, 415, 447, 422%, 682+
— — primigenius 93
— wusti 92, 417
Ellobius lutescens 141, 146*, 149, 154,
227;: 228, 230,,'234102473, 6"
— talpinus 149, 193, 194*, 473, 677*
Eostylocerus 41
Eotragus martinianus 31, 36*
Equidae 327—333
Equus 61, 71, 91, 100, 101, 129, 160, 163
— caballus 96, 97*, 112, 115*, 117, 119,
131. 4441, 150, 154%, 153, 10. 262)
1654201927435"! 43) 27a, 082"
Equus caballus gmelini 151, 433*, 434,
475, 684+
— — latipes 433*, 434
— hemionus 4154, 155, 188, 190,
NS, BPS aslo У ВМ
— hidruntinus 96, 97*, 117, 141, 150*,
151, 154, 683+
— przewalskii 433*, 434
— stenonis,59—61, 69, 70*, 74, 77, 100,
161, 432, 684+
— siissenbornensis 63, 67*, 77, 94, 129,
432, 684+
Erinaceidae 250
Erinaceus europaeus 122, 141—143, 153,
182, 183, 212, 219, 227, 228, 230, 250,
388, 466, 530, 547, 550, 645+
— —- rumanicus 645*
Eucladocerus 61, 63, 67*, 69, 70*, 71, 94
Eucladocerus pliotarandoides 60—62*
Felidae 270—283
Felis 109
— chaus 144, 155, 279, 469, 530, 550, 663+
— issiodorensis 52, 54%
— lybica 144, 144, 145*, 154, 225, 227,
469, 663+
— lynx 415%, 117, 144, 155, 175, 195,
277, 469, 550, 662+
— orientalis 405
— silvestris 99, 111%, 115, 144, 183, 214,
278, 547, 550, 662*
— — caucasicus 405
Gazella 39—44, 47, 52, 57, 63, 170%
— brevicornis 45
— gaudry 45
— subgutturosa 100, 152, 155, 188, 189%,
214, 215, 225, 354, 475, 530, 688+
Gerbillus 74
Giraffidae 41, 47, 52
Glis lis 183, 242, 473, 544, 547, 550,
66
Gulo gulo 122, 124*, 127, 266, 658+
Halitherium schinzi 26
Helicophora rotundocornis 45
Helladotherium 39
— gaudry 45
Hemiechinus auritus 141—143, 154, 182,
183, 191, 218, 219, 225, 230, 231, 254,
388, 466, 530, 533, 645+
Hipparion 41—43, 47, 52, 55*, 60, 69, 74
— crassum 69
— gracile 38, 39, 41, 43, 44, 47, 52, 55*
— mediterraneum 44
Hippopotamidae 31, 35*
Hippopotamus 160
Hyaena 31, 32*, 40*, 41, 59, 60
— eximia 39
— hyaena 155, 261—263, 469, 655+
— striata 129, 144
Hyaenidae 260—263
Hypsodontus miocenicus 31, 36*
Hystricidae 41
Hystrix 127, 130*, 681+
— hirsutirostris 117
— leucura 474, 681+
189*,
|
— vinogradovi 141, 146*, 149, 154, 681+
Iniops 26
— caucasicum 26
Insectivora 52, 123, 141, 233, 250—255,
388, 465, 476, 530, 533, 534, 547, 55
Ictitherium tauricum 44
Kubanochoerus robustus 31, 36*
Lagomorpha 52, 74, 123, 144, 233, 283 —
285, 405, 469, 473, 476, 530, 534, 547,
550, 692+
Lagurus lagurus 323, 473, 676+
Leporidae 283—285
Lepus 52, 53*, 74, 101, 664+
— europaeus 96, 99, 127, 149, 154, 175,
177, 179, 182, 183, 185, 186*, 191,
193, 195, 206, 208, 209*, 212, 214,
224%, 227, 228, 230, 231, 283—285,
473, 530, 547, 550, 664+
— — caucasicus 405
— — cyrensis 405, 534
ureevi 144, 146*, 664+
Listriodon 43
Lophiomeryx benarensis 27
Lutra lutra 144, 177, 469, 547, 550, 660*
Lutreola lutreola 144, 659*
— — turovi 404
— vison 691+
Macaca 127, 130*
Machairodus 47, 60
Marmota 127, 130*, 160, 664+
— baibacina 573, 692+
— bobac 179, 664+
Martes 108%, 109, 141%, 115%, 193, 402
— foina 117, 127, 144, 155, 224*, 225,
227, 267, 469, 550, 656+
— — nehringi 404
— martes 268—270, 469, 657+
Mastodontidae 71
Mastodon 31, 39, 44, 60
— arvernensis 59, 66, 69, см.
Anancus
— borsoni 48
— longirostris 39
— pentelici 39, 44
— tapiroides 48
Megaceros 103, 160, 161
— euryceros 63, 94, 96, 97*, 99, 109,
141, 152, 153, 346, 685+
— hibernicus 100
Meles 59, 660+
— meles 99, 415%, 117, 127, 144, 153, 160,
177, 179, 183, 193, 208, 209*, 222,
227, 469, 530, 547, 550, 660+
— — minor 141, 445%, 402,* 660+
— — urartuorum 660+
— maraghanus 660+
— polaki 660+
Mephitis mephitis 691+
Meriones 225, 228, 473, 676+
— blackleri 230, 305, 473, 534, 676+
— erythrourus 154, 218, 219, 473, 530,
533, 675*
также
813
— — intermedius 141, 146*, 414, 675+
— meriodianus 473, 675+
— persicus 227, 230, 231, 306, 473, 675+
— tamariscinus 473, 675+
— vinogradovi 473, 676+
Mesocricetus 47, 408
— auratus 112, 118, 119, 122, 124*, 127,
130%; 154, 182, 183. 185.1 195; 22%.
228, 230, 231, 303, 473, 673+:
— — nigriculus 186%, 191, 408
— — planicola 141, 146%, 408, 673
— — raddei 408, 873*
Micromeryx flourensianus 31
Micromys minutus 149, 182, 473, 672+
Microtinae 208
Microtus 411
— apscheronicus 141, 149, 413, 679+
arvalis 141, 149, 153, 182, 183, 185,
206, 208, 230, 231, 312, 414, 474, 550,
680+
— gudauricus 414
— штасгосгап!5 414
— mystacinus 414
— transcaucasicus 414
gud 127, 185, 317, 474, 678+
— gud 413
— lIghesicus 413
— nenjukovi 413
guentheri 314
majori 127, 182, 183, 208, 228, 315,
474, 547, 679+
nivalis 227, 228, 413, 320, 474, 678+
— dementievi 413
— loginovi 413
— satunini 413
roberti 115%, 180, 322, 474, 679+
socialis 141, 149, 154, 191, 218, 219,
227—231, 308—312, 415, 474, 530,
532—534, 544, 550, 680+
— — schidlovskii 309
Microzeuglodon 26
— caucasicum 26
Miniopterus schreibersii 230, 467, 650+
Muridae 71, 96, 127, 297—327
Mus 52, 671+
— musculus 141, 149, 153, 182, 183 185,
206, 208, 218, 219, 225, 230, 408, 473,
547, 566, 671+
abbotti 532, 550
formosovi 544, 671+
hortulanus 671+
musculus 530, 532—534
— tataricus 530, 532—534, 544, 550,
566, 671+
Mustela erminea 469, 658+
— filholi 74
— nivalis 127, 144, 155, 183, 185, 186%,
206, 208, 227, 230, 469, 530, 534,
547, 550, 659+
— — caucasica 122, 182, 228, 404
-- — dinniki 182, 404
Mustelidae 266—270
Myocastor coypus 525, 547, 692+
Myotis 390
467, 649+
lead а peak
|
i a a
— bechsteinii
— emarginatus 467, 649+
— mystacinus 467, 528, 534,
—- nattereri 467, 649+
— oxygnathus 227, 228, 467,
649+
649+
Necromites nestoris 69
Neomys 127, 389
— balcaricus 289
— daghestanicus 389
— fodiens 228, 389, 466, 646+
— leptodactylus 389
— schelkovikovi 389
Nyctalus leisleri 467, 651+
— noctula 467, 528, 651+
— siculus 467, 651+
Nyctereutes procyonoides 524, 691+
Ochotona 171, 227, 228, 473, 664+
— antiqua 52, 664+
— pusilla 664+
— rufescens 664+
Oiceros atropatanes 45
— boulei 45
— rothi 45
Ondatra zibethica 525, 692+
Orictolagus cuniculus 692+
Orycteropus sp. 31
— gaudryi 44
Otocolobus manul 469, 663+
Ovis 127, 195
— ammon 119, 127, 130%, 144, 152%,
154, 373, 690+
— яме 163, 195%, 225, 227, 373, 475,
— ophion 109
Palaeocricetus caucasicus 31, 674+
Palaeoloxodon 416
Palaeomastodon 32
Palaeoryx longicephalus 48
— pallasi 45
Panthera 63, 67*, 661+
— leo 144, 155, 270—274, 469, 6641+
— pardus 127, 130%, 144, 155, 171, 193,
194», 241, 275—277, 403, 469, 661+
— spelaea 99, 104, 115%, 117, 119, 127,
141, 144, 145%, 153, 270, 661+
— tigris 274, 469, 661+
Paracamelus 77
— gigas 69, 71*, 100, 685+
— ujalnikensis 685+
Parahippus 32
Paranchitherium karpinskii 31, 34+
Paratragocerus caucasicus 31, 36*
Parelephas —see_ Elephas
Perissodactyla 27, 31, 39, 41, 43, 44, 47,
52, 59, 60, 64, 63, 69, 74, 74, 123,
141, 233, 327—333, 430, 474, 476.
Phangare verve mammontoides 63, 415,
Phiomia 32
Phoca 28, 39, 47
— caspica 62, 69, 188, 215
— pontica 43
— vindobonensis 28
Phocidae 37
Phronetragus arknethensis 41
814
Pinnipedia 39, 43
Pitymys 315, 562, see Microtus majori
Platybelodon danovi 31, 33*
Plecotus auritus 467, 650+
Pliocervus 52
Primates 39, 44, 44, 233
Proboscidae 31, 39, 41, 44, 47, 48, 52,
59—63, 69, 71, 233, 415
Procapreolus 52, 55*, 687+
Procyon lotor 526, 550, 694+
Prometheomys 47, 411
— schaposchnicovi 122, 124%, 127, 323,
473, 677+
Propotamochoerus provincialie 52, 55*
Protoryx carolinae 45
Protragelaphus scozesi 45
Pseudalces 52
Putorius 74
— eversmanni 144, 183, 185, 469, 658+
— putorius 469. 658+
Rattus norvegicus 155, 182, 183, 185,
186*, 206, 212, 219, 299, 473, 530, 533,
547, 566, 670+
— rattus 208, 209%. 297, 473, 530, 547,
550, 670+
Rhinoceros 43, 60, 74, 127, 130°, 164
— antiquitatis 92
— binagadensis 141, 150, 151%, 154. 430,
431*, 683+
etruscus 61, 63, 71, 74, 160, 430, 683+
mercki 74, 92, 129, 164, 430, 683+
morgani 44
schleiermacheri 32
— tichorhinus 102, 430, 431*, 683+
Rhinocerotidae 38, 47, 52, 56°
Rhinolophidae 390
Rhinolophus blasii 467, 648+
— euryale 467, 648+
— ferrum-equinum 109; 467, 528. 648+
— hipposideros 467, 528, 648+
— mehelyi 467, 648+
Rhombomys opimus 187, 473, 675+
Rodentia 27, 31, 41, 52, 59, 60, 63, 71, 74,
123, 141, 233, 285—327, 405, 470, 473,
476, 530, 533, 534, 547, 550, 692+
Rupicapra гир!сарга 108*, 111*, 115%,
147, 118, 122, 193, 475
— — caucasica 202, 205, 241, 360, 688+
Saiga borealis 441%, 443
— prisca — see S. tatarica
— tatarica 96, 97*, 100, 357, 441*—443,
475, 687+, 692+
= АНИ 144, 152*, 153, 441*,
— — mongolica 442+
Schisotherium chu¢ua 27*, 27
Scirtopoda telum 473, 669+
Sciuridae 285—293
Sciurus anomalus 473, 550, 664+
— vulgaris 692+
— — altaicus 570
Semantor macrurus 69
Sicista betulina 473, 667+
— caucasica 96, 97*, 473, 667+
— subtilis 473, 667+
Sorex 250, 389
— araneus 185, 208, 466, 547, 550, 646+
— minutus 228, 389, 466, 646+
— raddei 389, 466, 646+
Spalax giganteus 187, 473, 669+
— leucodon 473, 669+
— microphtalmus 77, 182, 183, 185, 473,
669+
Spelaearctos rossicus 91
Stegodon 71
Steneofiber 52, 53%, 293, 665+
Strepsicerotini 63
Suidae 334—338
Suncus etruscus 466, 647°
Sus 41, 43, 47, 60
— apscheronicus
435, 436°, 684
— erymanthius 39, 44
— scrofa 96, 108*, 109, 111%, 116—118,
422, 127, 152, 175, 177, 183, 188, 193,
195, 202, 205, 208, 210, 222. 334, 435,
475, 530, 547, 550, 684+
— — attila 435, 436°
— tamanensis 63, 67%, 334, 435, 436*,
684°
141, 152%, 154, 334,
— vittatus 434
Tadarida taeniotis 467, 653+
Talpa 52, 53%, 127, 250, 388, 645+
— coeca 547
— caucasica 122, 124%, 182, 183, 186%,
251, 466, 645°
— — caucasica 389
— — ognevi 388, 389, 645*
— orientalis 251, 466, 645+
— — orientalis 389
— — talyschensis 388, 389
— — transcaucasica 388, 389
— caucasicus 63
Tapirus arvernensis 52, 55*
Tragelaphus 63
— hontom-schindleri 45
Тгаросегиз 39, 40%, 41, 44, 74
— leskevitschi 39, 41, 41*, 44
— rugosifron 45
Trogontherium cuvieri 59, 60, 63, 67*,
71, 665+
Tubulidentata 31, 44
Udabnopithecus garedziensis 41
Urmiabos azerbaidzanicus 45, 691+
Ursavus 31, 263
Ursidae 263—266
Ursus 60, 263
— arctos 109, 444, 447, 118, 144, 153,
183, 188, 189*, 208, 209*, 214, 222,
227, 264—266, 402*, 469, 550, 656*
— arctos binagadensis 141, 145*, 402%,
656+
— — caucasicus 122, 399, 401*, 402*
— — meridionalis 122, 402*
— arvernensis 52, 54*
815
kamiensis 400—402* Vormela peregusna 141, 145%, 154, 183,
karmalkiensis 401%, 402° 227, 469, 530, 657+
rossicus 263, 655° Vulpes 99
spelaeus 96, 99, 105, 108*—112, 115*, — corsac 141, 145%, 153, 259—260, 399°,
117—119, 122, 127, 130%, 264, 655+ 469, 655+
Vespertilio 390 — fulvus 694+
— bobrinskii 467, 652+ — khomenkoi 57, 391, 654+
— kiblii 467, 528, 533, 652+ — vulpes 108%, 109, 115%, 117, 122, 127,
— murinus 467, 652+ 144, 153, 179, 183, 188—190, 193,
— nathusii 467, 528, 652+ 206, 215, 222, 224°, 227, 258, 469,
— nilssonii 467, 653+ 530, 547, 550, 654+
— ognevi 467, 653+ — — alpherakyi 141, 145°, 392, 654+
— pipistrellus 467, 528, 534, 651+ — — alticola 392
— savii 467, 652+ — — kurdistanica 392
serotinus 467, 528, 533, 653+
Zeugiodon 26
1704 816
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