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Full text of "Mammals of the Soviet Union"

Mammals 

of the 
Soviet Union 




VG. Heptner 



A. A. Nasimovich 



A. G. Bannikov 




This volume of Mammals of the Soviet Union 
is devoted to a description of artiodactyls 
and perissodactyls found in the Soviet 
Union. These animals are of great scien- 
tific and economic interest; information 
on them, especially in recent Soviet 
literature, is voluminous. In recent years 
world literature on this subject has 
likewise been considerably enriched with 
new data. The abundance of information 
on ungulates explains the size of the 
present volume. 

All the characteristics of groups have 
been described according to a common 
scheme; deviations occur in a few cases, 
however. These characteristics are stated 
briefly and pertain to the group as a 
whole; they are not exclusive to species of 
the Soviet fauna. Diagnosis is based essen- 
tially on morphology Considerable atten- 
tion has been given to the composition 
of the group, its taxonomic relationships, 
and the position of Soviet forms in the 
world fauna. 

All species are described according to 
a common plan, altered only in the case 
of some extinct forms. In devising the 
scheme for descriptions of species not 
only the convenience of the reader was 
kept in mind, but the hope that gaps in 
our knowledge would become self-evident 
and stimulate further research. 

(from the Foreword) 




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SMITHSONIAN INSTITUTION 
WASHINGTON, D.C. 20560 

ASSISTANT SECRETARY FOR RESEARCH 



June 1990 



Dear Colleague: 

It is my pleasure to send you a copy of 
the English edition of Mammals of the Soviet 
Union , volume 1 (Artiodactyla and 
Perissodactyla) . This basic work, authored 
by Professor V.G. Heptner and colleagues, 
first appeared in 1961. It was translated 
into English by the Smithsonian Institution's 
Translation Publishing Program, and I served 
as scientific editor. I hope you will find 
the book useful to you in your work. It is 
anticipated that volume 2, part 2 will be 
published in the near future, followed by 
volume 2 , part 1 . I would be happy to hear 
from you if you believe that these subsequent 
volumes would be of interest to you. 

Sincerely, 




Robert S. Hoffmann 
Assistant Secretary 
for Research 



Mammals of the Soviet Union 

Volume I 



MAMMALS OF THE SOVIET UNION 
In Three Volumes 



Edited by 
V.G. Heptner and N.P. Naumov 



Vysshaya Shkola Publishers 
Moscow, 1961 



Mammals of the 
Soviet Union 

Volume I 
ARTIODACTYLA and PERISSODACTYLA 



V.G. Heptner, A. A. Nasimovich 
and A.G. Bannikov 



Scientific Editor 
Robert S. Hoffmann 




Smithsonian Institution Libraries 

and 

The National Science Foundation 

Washington, D.C. 

1988 



ТТ 76-52040 



Mlekopitayushchie Sovetskogo Soyuza 

In Three Volumes 

V.G. Heptner &: N.P. Naumov, editors 

Vysshaya Shkola Publishers 
Moscow, 1961 

Translator: P.M. Rao 

General Editor: Dr. V.S. Kothekar 

© 1988 Amerind Publishing Co. Pvt. Ltd., New Delhi 

Library of Congress Cataloging in Publication Data 

Geptner, V.G. (Vladimir Georgievich), 1901-1975. 
Mammals of the Soviet Union. 

Translation of: Mlekopitaiushchie Sovetskogo Soiuza. 

Contents: v. 1. Artiodactyla and Perissodactyla. 

Supt. of Docs, no.: SI I.2:Ar7/2 

1. Mammals — Soviet Union — Collected works. 
I. Nasimovich, A. A. II. Bannikov, Andrei Grigor'evich. 
III. Hoffmann, Robert S. IV. Title. 
QL728.S65G47 1986 599.0947 85-600144 

Translated and published for the Smithsonian Institution Libraries, pursuant 
to an agreement with the National Science Foundation, Washington, D.C., 
by Amerind Publishing Co. Pvt. Ltd., New Delhi, India 

Printed at Model Press, New Delhi, India 



Foreword to the English Edition 



Dr. Vladimir Georgievich Heptner, the senior editor and author of 
Mammals of the Soviet Union and Professor at Moscow State Univer- 
sity, died in Moscow on 5 July 1975, in his 75th year. Professor 
Heptner was born in Moscow in 1901. From the day he entered 
Moscow University as a graduate student in 1919, Professor Heptner's 
scientific and academic career was indissolubly linked with Moscow 
University's Faculty of Biology, and with its Zoological Museum. In 
1929, he became curator of mammals at the Museum and assistant 
professor; in 1934, he attained the rank of full professor. 

Heptner combined his intensive scientific work with continuous 
academic activity. Many Soviet biologists, zoogeographers, morphol- 
ogists, ecologists, and systematists, as well as mammalogists count 
themselves among his students, and his high esteem among his Soviet 
colleagues is manifested by their naming about 20 taxa of animals of 
different classes in his honor. Professor Heptner's scientific and peda- 
gogic activities made him a leader of Soviet zoology and one of the 
world's most renowned mammalogists. He was one of the oldest 
members of the Moscow Society of Naturalists and of the Geographi- 
cal Society of the USSR, and an honorary member of the All-Russian 
Society of Wildlife Conservation. In 1974, when the Theriological 
Society of the USSR was founded, he was made its vice-president. His 
scientific contributions were recognized in many foreign countries: 
he was an honorary member of the American Society of Mammalo- 
gists, the Gesellschaft Naturforschen der Freunde zu Berlin, the Deut- 
schen Gesellschaft fiir Saugetierkunde, and the Zoological Society of 
Czechoslovakia. He was also a member of the European Society of 
Mammal Protection, and of the International Union for Conserva- 
tion of Nature and Natural Resources. 

It is regrettable that during this long and productive career only a 
small number of non-Soviet mammalogists were fortunate enough to 
become personally acquainted with his keen mind and breadth of 
interests. Heptner is best known to American mammalogists for his 
publications in systematics and zoogeography. He dedicated many 
years of his life to the problems of zoogeography of the USSR, and of 
the Holarctic as a whole. In particular, his numerous studies of 
Middle and Central Asia and Asia Minor, such as his detailed investi- 
gations of Old World deserts, and his insights into the role of environ- 
ment in adaptation and differentiation of desert animals, are of great 



VI 



significance. His contribution to systematics of various mammalian 
groups is also large. One may note especially his work on systematics 
of Gerbillinae, a group in which he was especially interested. In the 
course of his systematic activity, he described about 70 new forms 
(subspecies, species, and subgenera) of mammals. Evolutionary prob- 
lems were of the greatest interest to Professor Heptner, especially 
those of microevolution and subspeciation. He was active not only as 
a scientist, but a popularizer in the USSR of the polytypic species 
concept. In this connection, he initiated the translation into Russian 
of such books as Ernst Mayr's Systematics and the Origin of Species 
(in 1947); Animal Species and Evolution (in 1968); Principles of 
Systematic Zoology (in 1971); and Populations, Species and Evolu- 
tion (in 1974); Mayr, l^insely andUsinger's Methods and Principles of 
Systematic Zoology (in 1956); and A. Cain's Animal Species and their 
Evolution (in 1958), all published under his editorship. Professor 
Heptner also served on editorial boards of various periodicals, includ- 
ing Zoologicheskii Zhurnal, Doklady Vysshei Shkoly, Okhota i 
okhotnichie khozyaistvo, Lynx, Das Pelzgewerbe, and Sdugetier- 
kundliche Mitteilungen. 

An excellent field naturalist, Heptner took part in many expedi- 
tions to all parts of the Soviet Union. Under his leadership field 
parties investigated such diverse areas as the Soviet Arctic, Middle 
Russia, the northern Caucasus Mountains, Transcaucasia, Turkme- 
niya, Uzbekistan, the Pamir, the Altai Mountains, and the Soviet Far 
East. His familiarity with the natural wealth of the USSR, and his 
appreciation of the problems attendant upon utilization of the coun- 
try's resources, led him to assume a pioneering role in wildlife conser- 
vation. At the same time, his broad approach permitted him to work 
effectively in "applied" fields such as medical zoology, pest control, 
fur harvest, and commercial hunting. 

Professor Heptner was the author or co-author of about 300 pub- 
lished works, among them such monographs as Mammals of the 
Middle Kopet Dagh and Adjacent Plains (1929), General Zoogeo- 
graphy (1963), Rodents of Middle Asia {19S6), Vertebrate Animals of 
Badkhyz (1956), and Harmful and Useful Mammals of the Protective 
Forest Zones (1950), the latter having been translated into German. 

His colleagues, both Soviet and foreign, feel his loss deeply — he 
was a man of great culture, integrity, fortitude, and generosity, 
always ready to help anyone with his words and deeds. (Modified 
from obituary in /. Mammal, 57 (2): 416-417, by Rossolimo and 
Hoffmann.) 

I first met Professor Heptner in May, 1963, two years after the 
publication of the first volume of wh. e regarded as his major life 



Vll 

work, Mammals of the Soviet Union. He considered it to be the 
successor, and supplement, to S.I. Ognev's Mammals of the USSR 
and Adjacent Countries, a work published in seven volumes between 
1928 and 1950. Heptner commenced his series with the ungulates, 
since Ognev had not lived to complete his task, and this was the 
major omission. The present volume is thus an invaluable extension 
of the coverage of Ognev's works. Heptner expressed great concern 
to me in 1963 that, while Ognev was now being translated into 
English (the first three volumes had just appeared in 1962, up to 
thirty-five years after their original publication), they were in many 
respects out of date. We discussed the desirability of fostering an 
English edition of his new series, and he expressed the hope that this 
might be done promptly. Unfortunately, Professor Heptner died in 
early summer of 1975, and so did not live to see either an English 
translation of his monograph, or even its completion in Russian; at 
the time of his death, two parts of volume two had appeared, and a 
third was posthumously published in 1976. Since then, responsibil- 
ity for the series has been assumed by Acad. Vladimir E. Sokolov, 
Director of the Institute of Evolutionary Animal Morphology and 
Ecology of the Academy of Sciences of the USSR, who assures me 
that additional volumes will appear in due course. 

The English translation project was initiated at my urging in 
late 1975 as a contribution to joint U.S.-U.S.S.R. studies on the 
evolution of Holarctic mammals sponsored by the Environmental 
Agreement between the two countries, and their Academies of Science. 
The Smithsonian Institution Libraries in cooperation with the 
National Science Foundation sponsored the translation of this as well 
as of hundreds of other scientific studies. The Smithsonian Libraries 
program, funded with Special Foreign Currency under the provisions 
of Public Law 480, represents an investment in the dissemination of 
knowledge to which the Smithsonian Institution is dedicated. The 
translation of volume one was ably completed by P.M. Rao of 
Amerind Publishing Co., New Delhi, India, under the general editor- 
ship of Dr. V.S. Kothekar. I subsequently read the translation in 
manuscript in order to check for scientific accuracy, and translitera- 
tion of proper names. 

Geographic names are generally transliterated directly, but a few 
exceptions were permitted {e.g. Moscow instead of Moskva, transla- 
tion rather than transliteration of certain modifiers of place names, 
such as Northern, rather than Severnaya, Dvina). Soviet administra- 
tive units are numerous, and the following equivalents were 
employed in translation: Krai, territory; oblasl', district; raion, 
region; guberniya (archaic), province. Also, in the original Russian 



VUi 

text, rivers, mountain ranges, and cities are often not explicitly identi- 
fied, the Soviet reader being presumed sufficiently familiar with the 
geography of the country to be able to understand from the context of 
the sentence what sort of place is referred to. Complicating the matter 
is the lack of articles as parts of speech in Russian. To assist the 
English reader, I have adopted the following conventions: if a river is 
referred to, an article precedes it; if a mountain range is referred to, it 
is translated as a plural; if a city is referred to, it is singular, and lacks 
the article. Examples are: the Ural (river); the Urals (mountains); 
Ural'sk (city). Geographic place names are also inflected in Russian, 
and these have been simplified by omitting transliteration of the 
inflected ending. For example, the Russian phrase v Yaroslavskoi г 
Kostromskoi oblastyakh is translated "in the Yaroslavl and Kostroma 
districts." In cases where the nominative form of the place name has 
an -sk ending this is, however, transliterated {e.g., Omsk); when a 
Russian "soft sign" is employed in a place name, this is transliterated 
as an apostrophe {e.g., Khar'kov). Because of the large number of 
place names in this volume, it was not possible to verify all of them, 
and some inconsistencies are likely to occur. I would appreciate it if 
readers would bring any errors they may notice to my attention. 

Robert S. Hoffmann 
Assistant Secretary for Research 
Smithsonian Institution 
Washington, D.C. 20560 



Foreword 



5 This volume of Mammals of the Soviet Union is devoted to a des- 
cription of artiodactyls and perissodactyls found in the Soviet 
Union. These animals are of great scientific and economic interest; 
information on them, especially in recent Soviet literature, is volum- 
inous. In recent years world literature on this subject has likewise 
been considerably enriched with new data. The abundance of infor- 
mation on ungulates (more than on other mammals) explains the 
size of the present volume. 

The format and contents of this book require a comment or two. 
All species of Artiodactyla and Perissodactyla which have ever inha- 
bited the Soviet Union, including those now extinct, are described. 
Total coverage provides a correct picture of present-day fauna and 
permits an evaluation of changes in it. Only rhinoceri, information 
on which is completely fantastic (see text), and the parabubal ante- 
lope (a species of Alcelaphinae; Vereshchagin, 1956) have been 
excluded; their occurrence in Russia in the historic past requires 
further confirmation. 

Simpson's (1945) system, the latest and most comprehensive, has 
been adopted as the basis for classification. This system covers the 
entire class down to genera and includes both present-day and fossil 
forms. True, it has some drawbacks and debatable points but their 
correction needs extensive investigation, which falls outside the pur- 
view of the present work. Hence only relatively insignificant devia- 
tions from Simpson's system have been incorporated and mainly 
concern the number and scope of generic groups. The latter, of 
course, at least with reference to recent fauna, ought to be delineated 
in greater detail. The changes introduced here have been defined in 
the text. 

The number of species of artiodactyls, only recently clarified, has 
been adopted from Haltenorth and Trenze (1956) with minor modi- 
fications (oxen, goat, and camel; V.G. Heptner). 

All the characteristics of groups have been described according to 
a common scheme; deviations occur in a few cases, however. These 
characteristics are stated briefly and pertain to the group as a whole; 
they are not exclusive to species of Soviet fauna. Diagnosis is based 
essentially on morphology. Considerable attention has been given to 
the composition of the group, its taxonomic relationships, and the 
position of Soviet forms in world fauna. Paleontological referen- 



ces are given, drawn primarily from Simpson's work (1945), but 
occasionally including other sources as well. 

All species are described according to a common plan, altered 
only in the case of some extinct forms. In devising the scheme for 
descriptions of species not only the convenience of the reader was 
kept in mind, but the hope that gaps in our knowledge would 
become self-evident and stimulate further research. 

In describing species geographic distribution and biology are 
given primary attention. Geographic ranges are presented in general 
terms (boundaries) based on a critical analysis of the factual material 
available. Stray occurrences outside the frontiers of the range are 
usually not included and only the main reference points of the range 
boundary indicated. 

Descriptions are based on "reconstructed" geographic ranges for 
the historic past, the dynamics of which have been traced insofar as 
possible. This approach evolved from the conviction that present- 
day ranges and modern faunal complexes cannot be understood 
without reconstruction of earlier ranges. Without such a historic 
perspective, present-day ranges often give an altogether distorted 
picture of the zoogeographic and ecological features of present-day 
species and the composition of extant fauna. Furthermore, a recon- 
structed range provides an excellent basis for undertaking practical 
measures (acclimatization and reacclimatization). Geographic 
ranges falling within the USSR have been detailed to the best of our 
ability but sometimes minor variations in time and space could not 
be traced. By and large the widest spread of species (in a given 
interval of time or in general) has been considered. 

In those cases where the geographic range underwent significant 
changes, beyond the magnitude of normal boundary variations, 
these have been traced and described as completely as possible. This 
phenomenon is extremely interesting both from theoretical and eco- 
nomic points of view. Only very general remarks are made concern- 
ing the geographic ranges in the prehistoric period and the 
Pleistocene. All the geographic range maps are original and were 
plotted from material collected for this publication. 

Geographic ranges falling outside the USSR, including recon- 
structed ones, are highly schematic and based on a large number of 
special studies and important compilations: Seton (1909-1910); 



'For further details on reconstructed geographic ranges, see V.G. Heptner's work: 
Dinamika areala nekotorykh mlekopitayushchikh i antropicheskii faktor (Dynamics 
of geographic range of some mammals and the human factor). Sb. I'oprosy Geografii, 
1959. 



XI 

Miller (1912); Lydekker (1913-1916); Sowerby (1923); Anthony 
(1928); Shortridge (1934); Philips (1935); Heim de Balzac (1936); 
Allen (1939, 1938-1940, 1942); Cabrera and Yepes (1940); Chasen 
(1940); Rode (1943-1944); Harper (1945); Simpson (1945); Anderson 
(1946); Tate (1947); Prater (1947); Hainard (1949); Roberts (1951); 
Ellerman and Morrison-Scott (1951); Burt and Grossenheider (1952); 
Morrison-Scott and Hayman (1953); Miller and Kellog( 1955); Halte- 
north and Trenze (1956); Van den Brink (1958); and others (for 
which references have been given in the text). 

The morphological characters of species are very briefly stated 
since detailed descriptions are available for most in Soviet literature 
(deer, cavicorns, sheep, goats, and others). Only species found 
within the Soviet Union are described; however, the taxonomic posi- 
tion of each has been assessed within the genus as a whole. Geogra- 
phic variation in the USSR has been revised or critically reviewed. 
Forms inhabiting the USSR have, by and large, been drawn from the 
most recent compilations of mammalian species in the Soviet 
Union. 

A strictly limited number of synonyms are included, based on the 
following principles. Only natural synonyms are mentioned. For 
genera synonyms are presented only for those for which the Soviet 
forms serve as the type genus, or if the generic name is important, or 
is sometimes used (or was formerly used), or could be applied to 
Soviet species. All names, current or otherwise, have been included 
among the synonyms of a species in the following cases: 1) name 
assigned from specimens of a species found in Soviet territory; 2) 
name given from specimens of a species belonging to other territo- 
ries, but important and used or formerly used but persisting for 
some Soviet forms; 3) name given for specimens belonging to 
other territories, not used for Soviet forms but possibly relevant for 
some; and 4) name of nominal form invariably given irrespective of 
its type locality (whether within or outside the USSR). Current 
names have not been separately identified anrang the synonyms of a 
species, but are included in the list of subspecies inhabiting the USSR 
where synonyms pertaining to a given subspecies are also indicated. 
Synonyms of subspecies occurring outside the USSR have been ex- 
cluded. Synonyms for categories above genus have not been mention- 
ed; neither have references to their description and nomenclature 
or, if mentioned, only those based on Simpson's system. Names 
given for domestic forms (horses, goats, etc.) have not been used here 
for wild species; the first name given to the wild form is used as its 
species name. Descriptions of species within an order commence 
with the least specialized one. 



xu 

The total number of mammalian species is approximately 3,500 
and the number of species in Soviet fauna about 300 (Heptner, 
1956). 

In the preparation of this book each author contributed as fol- 
lows: V.G. Heptner has written the introduction, characteristics of 
groups, including generic, keys for identification, all of the chapter 
devoted to aurochs*, yaks, and tarpans, and the sections "Diagno- 
sis," "Description," "Geographic Distribution," and "Geographic 
Variation" for all species except goitered gazelle. Mongolian 
gazelle, and saiga. He also processed the synonyms for all genera and 
species. A. A. Nasimovich has written the sections "Biology" and 
"Economic Importance" for all species of artiodactyls except swine, 
aurochs, yak, gazelle, antelope, and saiga. A.G. Bannikov compiled 
the descriptions, apart from synonyms and geographic variation 
(described by V.G. Heptner) of saiga, goitered and Mongolian 
gazelles, and the sections "Biology" and "Economic Importance" of 
hog and wild ass. The authorship of some individual sections has 
been acknowledged in the form of author initials at the end of the 
corresponding section or chapter. V.G. Heptner is responsible for 
the classification adopted for the majority of the species, subspecies, 
etc. 

Literature of 1956 and 1957 and partly 1958 has been included. 
From the works of 1959 only some articles were included during 
editorial revision. The bibliography contains only cited works; the 
literature actually perused was naturally more extensive. Works 
referred to in giving synonyms of species and genera and those not 
used subsequently in the text have not been included in the 
bibliography. 

In the compilation of this monograph the authors naturally 
included in the text (in some cases acknowledged) their unpublished 
material. Moreover, a significant amount of unpublished data fur- 
nished by various authors has been included, namely: K.G. Abramov 
(Ussuri); Yu.V. Averin (Kishinev); I.I. Barabash-Nikiforov (Voro- 
nezh); O.P. Bogdanov (Tashkent); G.F. Bromlei (Vladivostok); I.V. 
Zharkov (Voronezh preserve); M.A. Zablotskii (Prioksko-Terras pre- 
serve); Yu.A. Isakov (Moscow); F.L. Ivanauskas (Kaunas); P.F. Kaz- 
nevskii (Voronezh preserve); S.G. Kalugin (Caucasian preserve); 
V.P. Kostin (Tashkent); V.A. Kotov (Caucasian preserve); L.S. 
Lebedeva (Moscow); A.N. Leont'ev (Chita); R.N. Meklenburtsev 
(Tashkent); K.D. Mamisashvili (Lagoda preserve); P.A. Merttsem 

*Some confusion exists in ttie Russian original with regard to turs (mountain 
goats) and aurochs — General Editor. 



XUl 

(Caucasian preserve); B.M. Petrov (Tashkent); A.S. and I.F. Popkov 
(Zakatal'sk preserve); P.P. Tarasov (Przheval'sk); V.V. Timofeev 
(Irkutsk); O.I. Semenov-Tyan-Shanskii (Lapland preserve); V.N. 
Skalon (Irkutsk); A.V. Fedosov (Bryansk); V.I. Chernyshev (Stali- 
nabad); F.D. Shaposhnikov (Tyumen); P.B. Yurgenson (Moscow); 
and several others (acknowledged at the appropriate place). 

Workers in almost all of the regional game inspectorates to 
whom requests were sent for information about the distribution of 
various species responded readily. Extremely informative material 
was furnished by comrades Starodubchenko (Kur); Lebedov 
(Stalingrad); Mikhaylovka (Saratov); Zakharov (Murmansk); Gusev 
(Rostov-on-Don); Khmelevskii (Orenburg); Sukhikh (Belgorod); 
Ivanov (Yakutia); Kuz'min and Fofanov (Novosibirsk); Bel'skii (Vla- 
divostok); Anashkin (Ulan Ude); Samsonov (Blagoveshchensk); 
Sysoev (Khabarovsk); and Zimin (Chita). Some material was provided 
by N.V. Eliseev, Director of the Central Board of Hunting and Pre- 
serves, Russian Soviet Federated Socialist Republic. 

Valuable unpublished material on distribution — references, 
maps, manuscripts, etc. — was furnished by V.Ya. Parovshchikov 
(Arkhangel'sk); K.A. Tatarinov (L'vov); A. P. Korneev (Kiev); and 
A. A. Sludskii (Alma-Ata); and especially by N.K. Vereshchagin 
(Leningrad) and S.V. Kirikov (Moscow). The authors express their 
deep gratitude to each and everyone of them, as well as to S.P. 
Naumov and S.S. Turov who reviewed the manuscript and offered 
several suggestions. 

The authors are also extremely indebted to V.N. Aleksandrov, 
V.A. Kotov, E.P. Knorr, G. Levenshtein, K.T. Yurlov, P.A. 
Yanushko, O.I. Semenov-Tyan-Shanskii, Yu.V. Averin, V.D. Treus, 
and several others whose names have been given at appropriate places. 
The authors are grateful for various illustrative material to H. Heck, 
Director of Munich Zoological Garden, who furnished a photo- 
graph of his reconstructed auroch and tarpan. V.A. Dolgov and V.N. 
Orlov rendered invaluable assistance in the preparation of the 
manuscript for the printer. 

The authors received much cooperation in various forms from 
the Zoological Museum of Moscow University; the present volume is 
largely based on material provided by the university. 

In a work of such magnitude and extensive literary coverage, 
errors and omissions in information are inevitable. This is particu- 
larly so for data on distribution. How vague our knowledge still is, 

"To differentiate references to literature, references to personal communications 
have been identified by including the initials of the author and omitting the year. 



is evident in the distribution of argali sheep in Turkmenia, 
Gissar, and Zerevshan mountains. The authors request all readers, 
especially folklorists, hunters, and other specialists to send their 
comments, corrections, and supplementary information to Prof. Vla- 
dimir Georgievich Heptner, Zoological Museum, Moscow Univer- 
sity, Hertsen Street, 6, Moscow, K-9. 



Table of Contents* 

FOREWORD TO THE ENGLISH EDITION v 

FOREWORD ix 

CLASSIFICATION OF CLASS MAMMALIA xxv 

KEY FOR IDENTIFYING ORDERS OF MAMMALS xxvi 



PART I. ORDER OF EVEN -TOED UNGULATES 
ORDER ARTIODACTYLA OWEN, 1848 

Superorder Paraxonia Marsh, 1884 3 

Order Artiodactyla Owen, 1848 3 

Suborder Suiformes Jaeckel, 1911 11 

Infraorder Suina Gray, 1868 15 

Family Suidae Gray, 1821 (Swine) 15 

Boar, or Wild Swine: Genus Sus 

Linnaeus, 1758 16 

1. Sus (Sus) scrofa Linnaeus, 1758 19 

Diagnosis 20 

Description 20 

Taxonomy 25 

Geographic Distribution 26 

Geographic Variation 43 

Biology 45 

Economic Importance 78 

Suborder Tylopoda Illiger, 1811 83 

Suborder Ruminantia Scopoli, 1777 85 

Infraorder Pecora Linnaeus, 1758 91 

Superfamily Cervoidea Simpson, 1931 91 

Family Cervidae Gray, 1821 91 



*Pages 773-776 in Russian original. The English table of contents is not a literal 
translation. — 



Subfamily Moschinae Zittel, 1893 (Musk 

Deer) 98 
Genus of Musk Deer: Genus Moschus 

Linnaeus, 1758 100 

2. Moschus moschiferus Linnaeus, 

1758 101 

Diagnosis 101 

Description 101 

Taxonomy 105 

Geographic Distribution 105 

Geographic Variation 111 

Biology 112 

Economic Importance 122 

True or Plesiometacarpal Deer 124 

Subfamily Cervinae Baird, 1857 124 

Genus of True Deer: Genus Cervus 

Linnaeus, 1758 126 

3. Spotted [sika] deer: Cervus 

(Cervus) nippon Temmink, 1838 131 

Diagnosis 132 

Description 132 

Taxonomy 136 

Geographic Distribution 136 

Geographic Variation 141 

Biology 141 

Economic Importance 156 

4. True, Wellborn [Red] Deer: ^ 
Cervus (Cervus) elaphus Linnaeus, 

1758 159 

Diagnosis 161 

Description 162 

Taxonomy 167 

Geographic Distribution 167 

Geographic Variation 185 

Biology 190 

Economic Importance 230 
Subfamily of Elk, or Telometacarpal Deer: 

Subfamily Odocoileinae Pocock, 1923 235 



XVll 



Genus of Roe Deer: Genus Capreolus 




Gray, 1821 


238 


5. Roe Deer: Capreolus capreolus 




Linnaeus, 1758 


243 


Diagnosis 


244 


Description 


244 


Taxonomy 


249 


Geographic Distribution 


249 


Geographic Variation 


275 


Biology 


280 


Economic Importance 


299 


Genus of Elk: Genus Alces Gray 1821 


302 


6. Elk, Sokhatyi: Alces alces 




Linnaeus, 1758 


307 


Diagnosis 


308 


Description 


309 


Taxonomy 


314 


Geographic Distribution 


314 


Geographic Variation 


358 


Biology 


363 


Economic Importance 


417 


Genus of Reindeer: Genus Rangifer 




H. Smith, 1827 


425 


7. Reindeer: Rangifer tarandus 




Linnaeus, 1758 


430 


Diagnosis 


432 


Description 


432 


Taxonomy 


438 


Geographic Distribution 


438 


Geographic Variation 


458 


Biology 


463 


Economic Importance 


512 


Superfamily of Hollow-horned Ungulates: 




Superfamilia Bovoidea Simpson, 1931 


519 


Family of Hollow-horned Ungulates, or 




Catde: Familia Bovidae Gray, 1821 


519 


Subfamily Bovinae Gill, 1872 (Catde) 


532 



Genus of Aurochs, or True Cattle: 

Genus Bos Linnaeus, 1758 (Cattle) 537 

8. Aurochs, Primitive Cattle: Bos 

(Bos) primigenius Bojanus, 1827 539 

9. Yak: Bos (Poephagus) mutus 
Przevalski, 1883 549 

Genus of Zubr [ = Wisent]: Genus Bison 

H. Smith, 1827 554 

10. Wisent: Bison bonasus Linnaeus, 

1758 557 

Diagnosis 558 

Description 558 

Taxonomy 560 

Geographic Distribution 561 

Geographic Variation 571 

Biology , 573 

Economic Importance 598 

Subfamily of Gazelles: Subfamily 

Antilopinae Baird, 1857 599 
Genus of Gazelles: Genus Gazella 

Blainville, 1816 604 

11. Dzheiran: Gazella subgutturosa 
Gueldenstaedt, 1780 608 
Diagnosis 608 
Description 608 
Taxonomy 610 
Geographic Distribution 610 
Geographic Variation 617 
Biology 617 
Economic Importance 632 

Genus of Dzeren, or Mongolian 
Antelope (Gazelle): Genus Procapra 

Hodgson, 1846 634 

12. Mongolian Gazelle, or Dzeren: 
Procapra gutturosa Pallas, 1777 636 
Diagnosis 637 
Description 637 
Taxonomy 639 
Geographic Distribution 639 



Geographic Variation 642 

Biology 642 

Economic Importance 652 

Subfamily of Goats: Subfamily Caprinae 

Gill, 1872 653 

Genus of Saiga: Genus Saiga Gray, 

1843 659 

13. Saigak, Saiga: Saiga tatarica 
Linnaeus, 1766 662 
Diagnosis 662 
Description 662 
Taxonomy 666 
Geographic Distribution 666 
Geographic Variation 671 
Biology 672 
Economic Importance 696 

Genus of Gorals: Genus Nemorhaedus 

H. Smith, 1827 700 

14. Goral: Nemorhaedus goral 
Hardwicke, 1825 704 
Diagnosis 705 
Description 705 
Taxonomy 707 
Geographic Distribution 707 
Geographic Variation 711 
Biology 713 
Economic Importance 719 

Genus of Chamois: Genus Rupicapra 

Blainville, 1816 (Chamois) 720 

15. Chamois, Serna, Black Goat: 
Rupicapra rupicapra Linnaeus, 

1758 723 

Diagnosis 724 

Description 724 

Taxonomy 727 

Geographic Distribution 728 

Geographic Variation - 732 

Biology 734 





Economic Importance 




746 


Genus of Mountain, or Rock, Goats: 




Genus Capra Linnaeus, 1758 




748 


16. 


Bearded or Bezoar Goat: 


Capra 






(Capra) aegagrus Erxleben, 


1777 


756 




Diagnosis 




757 




Description 




757 




Taxonomy 




760 




Geographic Distribution 




761 




Geographic Variation 




766 




Biology 




768 




Economic Importance 




773 


17. 


Siberian Mountain or Rock Goat 
[Siberian Ibex]: Capra (Capra) 






sibirica Pallas, 1776 




774 




Diagnosis 




775 




Description 




775 




Taxonomy 




784 




Geographic Distribution 




785 




Geographic Variation 




795 




Biology 




798 




Economic Importance 




814 


18. 


Kuban Tur: Capra (Capra) 








caucasica Gueldenstaedt and 






Pallas, 1783 




816 




Diagnosis 




817 




Description 




817 




Taxonomy 




822 




Geographic Distribution 




823 




Geographic Variation 




825 




Biology 




826 


19. 


Dagestan Tur: Capra (Capra) 






cylindricornis Blyth, 1841 




826 




Diagnosis 




826 




Description 




826 




Taxonomy 




831 




Geographic Distribution 




833 




Geographic Variation 




837 



XXI 



Biology 838 

Economic Importance 857 

20. Screw-horned Goat or Markhor: 
Capra (Orthaegoceros) falconeri 
Wagner, 1839 859 

Diagnosis 859 

Description 860 

Taxonomy 863 

Geographic Distribution 864 

Geographic Variation 868 

Biology 870 

Economic Importance 874 

Genus of Mountain, or Rock, Sheep: 
Genus Ovis Linnaeus, 1753 874 

21. Mountain Sheep, Arkhar: Ovis 

ammon Linnaeus, 1758 881 

Diagnosis 884 

Description 884 

Taxonomy gQ2 

Geographic Distribution 903 

Geographic Variation 919 

Biology 929 

Economic Importance 952 
22. Snow Sheep, Bighorn Sheep, 
Chubuk: Ovis canadensis 

Shaw, 1804 954 

Diagnosis 955 

Description 955 

Taxonomy 959 

Geographic Distribution 959 

Geographic Variation 965 

Biology 969 

Economic Importance 978 



SUPPLEMENT TO ORDER OF EVEN -TOED 
UNGULATES 



981 



ARTIODACTYLA 981 

Deer Family 981 

Family Cervidae Gray, 1821 981 

Genus of True Deer: Genus Cervus 

Linnaeus, 1758 981 

23. Fallow Deer: Cervus (Dama) 

dama Linnaeus, 1758 981 

Diagnosis 981 

Description 981 

Taxonomy 984 

Geographic Distribution 985 

Geographic Variation 989 

Biology 990 

Economic Importance 994 



PART 11. ORDER OF ODD -TOED UNGULATES 
ORDER PERISSODACTYLA OWEN, 1884 



Superorder Mesaxonia Marsh, 1884 


997 


Order Perissodactyla Owen, 1848 


997 


Suborder of Horses: Suborder Hippomorpha 




Wood, 1937 


1005 


Superfamily Equoidea Hay, 1902 


1005 


Family Equidae Gray, 1821 


1005 


Subfamily Equinae Steinmann and 




Doderlein, 1890 


1009 


Genus of Horses: Genus Equus 




Linnaeus, 1758 


1009 


1. Kulan: Equus (Equus) hemionus 




Pallas, 1775 


1011 


Diagnosis 


1012 


Description 


1012 


Taxonomy 


1014 


Geographic Distribution 


1015 


Geographic Variation 


1023 


Biology 


1025 


Economic Importance 


1036 



XXIU 

2. Tarpan: Equus (Equus) przewalskii 

Poljakov, 1881 1037 

Diagnosis 1039 

Description 1039 

Taxonomy 1041 

Geographic Distribution 1041 

Geographic Variation 1046 

Biology 1051 

Economic Importance 1055 

LITERATURE CITED 1059 

INDEX OF LATIN NAMES OF TAXA* 1135 



*The Russian original also included an Index of Russian Names of Taxa, omitted 
here. 



Classification of Class Mammalia 



The main taxonomic subdivisions (up to orders) adopted in the 
present work are based on the monograph on taxonomy written by 
G.G. Simpson (1945), who made extensive use of paleontological 
data. It is very similar to the system developed by M. Weber (1928), 
which, in its time, was well founded on anatomical data of recent 
forms. The orders acknowledged by Simpson have long been recog- 
nized and their grouping as suggested by him into higher groups 
gives rise to no serious objections. 

Of late a tendency has developed for greater fragmentation of 
orders. It has been proposed that marsupials be divided into three 
orders (Polyprotodontia, Caenolestoidea, and Diprotodontia), ceta- 
ceans into two orders (toothless whales — Mystacoceti, and toothed 
whales — Odontoceti), primates into three orders (Prosimiae, Simiae, 
and Tarsioidea), and so on. All these suggestions, however, are as yet 
not well founded. Moreover, among the morphology and paleontol- 
ogy oriented mammalian systematists such a tendency to extreme 
division is not acceptable. We believe an order is primarily an 
integrating and not a differentiating concept. Otherwise the same 
situation will inevitably arise in mammalogy as has already arisen 
in ornithological taxonomy wherein the concept of an order has 
largely lost its meaning; as a result the scientific aspect of vertebrate 
systematics has suffered greatly. 

Only one change has been made in the orders under con- 
sideration — pinnipeds are treated as an independent order and not 
as a suborder of Carnivora. Based on the classification of modern 
mammals these two groups separate naturally, i.e., they are no less 
distinctive than some other orders. 

Based on already established views Lagomorpha are treated as a 
separate order (Weber considered them only a suborder of rodents). 
Thus, instead of the 18 orders of Simpson, 19 are recognized here. Of 
these, 10 (52.6%) are represented in Soviet fauna and one (Steller's 
sea cow) is now extinct. 

The classification adopted for the class is depicted below. Orders 
present in the Soviet fauna are asterisked (V.H.). 



XXVI 



10 



CLASS MAMMALIA 



Subclass PROTOTHERIA 

Subclass THERIA 

Infraclass METATHERIA 
Infraclass EUTHERIA 



Cohort UNGUICULATA 



Order MONOTREMATA 
Order MARSUPIALIA 

"*Order INSECTIVORA 
Order DERMOPTERA 

*Order CHIROPTERA 
Order PRIMATES 
Order EDENTATA 
Order PHOLIDOTA 



Cohort GLIRES 



*Order LAGOMORPHA 
*Order RODENTIA 



Cohort MUTICA 



*Order CETACEA 



Cohort 
FERUNGULATA 



Superorder 

FERAE 

Superorder 



*Order CARNIVORA 
*Order PINNIPEDIA 
Order 



PROTUNGULATA TUBULIDENTATA 



Superorder 

paenungulata" 



Superorder 
MESAXONIA 

Superorder 
PARAXONIA 



Order PROBOSQDEA 
Order HYRACOIDEA 

!Order SIRENIA 

*Order 
PERISSODACTYLA 

*Order 
ARTIODACTYLA 



11 



Key for identifying Orders of Mammals 



1 ( 2). Hind hmbs absent. Body fishhke with large bilobate caudal 

fluke set horizontally CETACEA 

2 ( 1). Hind limbs present. Body not fishlike; tail, if present, not in 

form of bilobate fluke. 
3(4). Forelimbs in form of leathery wings . . CHIROPTERA 
4( 3). Forelimbs of different structure. 



XXVll 

5 ( б). Fore- and hind-limbs very short, paddle-shaped, and in the 
form of fins, i.e., all digits right up to very tips enclosed in a 
common skin PINNIPEDIA 

6( 5). Fore- and hind-limbs of different structure, not in form of 
fins. 

7(10). Hooves on feet. 

8 ( 9). Only one hoof on each limb .... PERISSODACTYLA 

9( 8). Two large hooves and two small ones set above them occur 
on each limb ARTIODACTYLA' 

10 ( 7). Hooves absent on legs (claws present). 

11 (14). Diastema occurs between large, chisellike incisors and molars; 

its length not less than length of entire row of molars on 
corresponding jaw. 

12 (13). Two incisors on upper jaw RODENTIA 

13 (12). Four incisors on upper jaw; small blunt one occurs behind 

each of two large sharp ones LAGOMORPHA 

14 (11). Diastema between incisors and molars absent or much smaller 

than length of molar row. Canines present. 

15 (16). Anterior portion of snout extends into well-developed small 

proboscis. Anteriormost tooth on each jaw or only on upper 
jaw much larger than adjacent tooth.^ . . . INSECTIVORA 
16(15). Anterior portion of snout does not form proboscis. Anterior- 
most tooth on each jaw not larger than adjacent one . . . 
CARNIVORA^ 

(V.H.) 



Structure of limbs differs in camels. Eacfi limb ends in two broad callused pads 
and true hooves absent; replaced with two very broad claws. Camels are not included 
in this key because they are domesticated animals. 

"If tooth relation differs, proboscis always present and .anteriormost tooth 
extremely short. Wrist very broad with huge claws and set on edge — inner surface 
backward (moles). 

Skulls of extinct Steller's cow, a representative of the order of sea cows or Sirenia, 
have been found on the coast of Commander Islands. They are distinguished by the 
absence of teeth on either the upper or lower jaw over a length of about 60 cm. 



PART I 



ORDER OF EVEN-TOED UNGULATES 

Order Artiodactyla Owen, 1848 



Superorder PARAXONIA Marsh, 1884 
Order ARTIODACTYLA Owen, 1848 



15 Artiodactyla is one of the most specialized orders of terrestrial mam- 
mals and includes large, medium, and rarely, small-sized species, 
most of which are light in build and only a few heavy and bulky. 

The digits on the fore- and hind limbs generally number four, 
but in some cases two or four in the fore-limbs and three in the hind 
limbs. Correspondingly, but to varying degrees, other metapodial 
elements also undergo reduction. The limb axis invariably passes 
between digits III and IV (paraxonic), which are better developed 
than the other digits and carry the entire body weight in most cases. 
As a result of the great development (elongation) of the axial portion 
of the metapodium, the lateral digits are not only smaller than the 
medial but also set above them. The terminal phalanges of the digits 
are hooved and asymmetrical in shape. The terminal phalanges of 
the medial digits are greatly enlarged. Some have an enlarged toe- 
nail instead of a hoof. 

As a result of adaptation to fast running, all the limb sections in 
most of these animals are highly elongated. Simultaneously, diver- 
sity of movement (pronation and supination) was lost and confined 
to movement of the entire limb and bending and unbending of its 
component parts in a sagittal plane. Both the clavicle and the third 
trochanter of the femur absent. In most animals of this order the 
ulna is reduced and fused with the radius. Fibula either not fully 
developed or fused with the tibia. The astragalus has two (upper and 
lower) articular surfaces. There are usually 19 thoracicolumbar ver- 
tebrae. The tail in most animals is greatly reduced and either almost 
invisible externally or barely so; in other animals (oxen) it is quite 
long. 

As a result of the elongation of teeth and the tooth row, the skull 
usually has a protruding facial portion, sometimes elongated in the 
form of a wedge; a tendency toward pneumatization of individual 
bones is seen to some extent. The orbit is separated from the tem- 
poral fossa by the fused postorbital processes of the frontal and jugal 
bones, or these processes are not fused and a gap occurs between 
them. The lacrimal is well developed and its facial and orbital por- 
tions equal in size or the former greatly reduced. In some animals a 

16 well-developed ethmoid pit occurs before the lacrimal. The alisphe- 



noid canal is not developed. In most species horns are present in one 
or both sexes. With few exceptions/ the horns are a single pair and 
set on a projection of the frontal. The horns are bony and renewed 
annually or permanent^ consisting of horny material around a bony 

core. 

Dentition is heterodont and diphyodont. The complete dental 

formula is: 

i— c— Dm— m— = 44. 
3 ' 1 ' 4 3 

However, a great reduction in teeth is seen in most artiodactyls; the 
upper incisors, canine, and first premolar are absent, and the lower 
canine generally incisiform. The diastema is usually distinct, the 
tooth row extended, and the molars bluntly tubular (bunodont in a 
few) or selenodont, or brachyodont more often hypsodont, and 
sometimes prismatic. 

The stomach in most artiodactyls is complex (ruminant) and 
consists of several chambers; rarely, it is simple. The cecum varies in 
design or is even absent. In most species distinct aggregations of 
cutaneous glands are present. The hair coat generally consists of 
normal hair, usually very coarse, but is sometimes bristly or almost 
absent. Inguinal teats number one or two pairs commonly. In some 
forms (pigs), however, five or six pairs of inguinal and abdominal 
teats are present. The scrotum is usually well developed and absent 
only exceptionally (Hippopotamus and Sus) (testes here lie under 
the skin). The uterus is bicornuate and the placenta diffuse or poly- 
cotyledonary. Sexual dimorphism is often well marked and in some 
animals associated with season or age. 

In structure, external appearance, size, and biological character- 
istics, artiodactyls are extremely diverse and represented by well- 
known types such as pigs, hippopotami, camels, llamas, mouse-deer, 
musk deer, giraffes, oxen, goats, sheep, and the highly diverse group 
"antelopes". The largest forms (giraffes) have a body length of 
almost 5 m, overall height of up to 6 m, height at withers up to 3 m 
or slightly more (3.3 m) or (hippopotami) a body length of up to 4.5 
m, height at withers up to 1.5 m, and weight up to 2,650 kg. The 
smallest forms (Neotragus) have a height at withers of 25 cm, total 



^Tetraceros has four typical horns. Giraffa has two, three, or five horns set not 
only on the frontal but also on the parietal and nasal bones; no bony or horny portion 
is exposed in them. 

^Antilocapra (American pronghorn) is an exception. 



body length up to 52 cm, and weight 2 to 3 kg. 

Most artiodactyls are swift runners and some are well adapted to 
scaling cliffs. As a result, most are slender in general appearance and 
have long legs. They live in diverse environments — forests of all 
types (from humid tropical to elfin woodlands), shrub growths, 
steppes, deserts, tundras, marshes, and reed bushes at all altitudes 
right up to the tops of extremely high mountains. One genus (Hip- 
popotamus) leads an amphibious mode of life. 

These animals usually live in herds or in small groups and are 
rarely found singly. They do not construct shelters nor maintain 
permanent shelters. Many species perform seasonal or irregular 
migrations, often over considerable distances. They are predomi- 
nantly polygamous, and usually bear one or two offspring, rarely 
four, five, or seven, and in a few species up to ten and perhaps 
twelve. The young are highly developed at birth and soon able to 
run. Males do not participate in the care of the offspring. 

Most artiodactyls are strict herbivores and eat mainly grassy vege- 
tation and rarely shrubs or trees and more rarely aquatic vegetation. 
A few species exhibit a euryphagous tendency and consume vegeta- 
tive as well as animal food — mainly terrestrial and soil invertebrates 
and sometimes small vertebrates. 

Artiodactyls are characterized by extremely wide distribution. 
They are found in South America (from Tierra del Fuego), through- 
out all of North America, including the Arctic archipelago and 
18 Greenland (of the Aleutians evidently only Unimak), all of Africa, 
Madagascar, and Eurasia, in the north including Spitsbergen 
(absent in Iceland), Novaya Zemlya, the southern island of Sever- 
naya Zemlya, and New Siberian Islands (absent on Wrangel Island 
and on all Kuril Islands except probably Shumsha). In the south, the 
geographic range covers Ceylon, Nicobar and Andaman Islands, 
and the entire island region between Asia and Australia in the south, 
including Little Sunda Islands, Timor, Aru and Kai archipelagos. 
New Guinea, and Louisiade and Solomon Islands (reconstructed 
range). 

In diversity of form and biological types, geographic distribu- 
tion, and number of species and population size artiodactyls consti- 
tute the most prosperous group of ungulates. 

Modern artiodactyls have been grouped with perissodactyls (and 
placed in the same superorder). In reality these two groups are mor- 
phologically widely separated, and as far as can be judged from 

'Sambar on Guam Island (Mariana Islands) is an acclimatized Philippine species 

(Rusa philippinensis). 



paleontological material, cannot be regarded as closely associated 
phylogenetically. The phylogenetic relationship of artiodactyls 
with perissodactyls is not much closer than their relationship with 
carnivores, and even less than the relationship between hyraxes, sir- 
enians, and elephants. In the latest taxonomic systems the separa- 
tion of artiodactyls and perissodactyls is emphasized by assigning to 
each group the rank of superorder (Paraxonia and Mesaxonia). 
They are linked only within the cohort Ferungulata, which also 
includes carnivores, pinnipeds, hyraxes, elephants, and sirenians 
(Simpson, 1945). Differences in their parasitic fauna also underscore 
the distinct segregation of artiodactyls and perissodactyls. Several 
features of similarity between the two orders evidently arose through 
parallel evolution associated with adaptation to swift running and 
feeding on coarse, partly dry, herbaceous food. 

Remains of ungulates have not been found in Cretaceous forma- 
tions. Fairly diverse and widely-distributed ungulate faunas of the 
extinct orders Condylarthra, Amblypoda, and Notoungulata have 
been found at the commencement of the Tertiary period (Paleocene). 
Evidently ungulates developed from primitive placentals in the 
Upper Mesozoic. Artiodactyls appear in Lower Eocene formations 
of North America and Europe. One has to search among Condylar- 
thra for ancestors of artiodactyls as for some other groups of recent 
ungulates. Condylarthra are closely associated with primitive carni- 
vores (extinct Creodonta). 

Perissodactyls were already well-developed in the Lower Eocene. 
Both groups appeared simultaneously, probably from the same 
stock. In Lower Tertiary horizons artiodactyls were fewer numeri- 
cally than perissodactyls. This ratio gradually changed later. Artio- 
dactyls, mainly Ruminantia, began to increase numerically and in 
the Upper Tertiary period formed the predominant group of ungu- 
lates. This ratio persists even to the present day. 

The classification of artiodactyls presents considerable difficul- 
ties, mainly because of the large number and diversity of forms. The 
problem is further complicated when fossil forms are also taken into 
consideration. Hence the systems of classification proposed by differ- 
ent authors vary greatly. It is rational to adopt three suborders — 
Suiformes, Tylopoda, and Ruminantia. All fossil forms also fit into 
these suborders. 

The order Artiodactyla comprises 419 genera included in 25 fami- 
lies, of which 16 are extinct and 9 extant; extinct genera number 333 
and extant 86 (Simpson, 1945). 

''in evaluating these and similar figures one must bear in mind that comparing 



19 To establish accurately the number of extant species in an order 
is difficult because of our poor understanding of the taxonomy of 
species. Probably they number about 150 (according to Haltenorth 
and Trenze, 1956). Thus they number more than all the remaining 
ungulates taken together and comprise about 4.5% of all mammal- 
ian species. On the average each genus contains 1.6 species and every 
family about 17. The largest number and diversity of species is seen 
in Africa (72 species) and Asia (65 species); the corresponding figures 
for North America, South America, and Europe are 12, 15, and 11 
species. 

In most parts of the world wild artiodactyls are of definite and 
sometimes great economic importance as game animals, providing 
meat, hide, and other products for making consumer goods, and raw 
material for pharmaceutical preparations. They are valued every- 
where as sport objects. Many species have greatly reduced in 
numbers and their range has shrunk; other have been totally de- 
stroyed; and several are now under total or partial protection. Artio- 
dactyls are one of the important animal groups protected under 
conservation programs in nature preserves throughout the world. 
Some species have been acclimatized while others serve as controlled 
game animals. 

Some artiodactyls are responsible for the spread of diseases, 
including protozoans (trypanosomiasis), among man and domestic 
animals. Consequently they are destroyed in some areas (tropical 
Africa). Together with perissodactyls they include the most impor- 
tant domesticated animals. Some species are both wild and domesti- 
cated or semidomesticated. 

Artiodactyl fauna of the Soviet Union comprise (excluding 
domesticated animals and fallow deer) a total of 19 species, about 
1.5% of the species of the order and about 6.3% of the mammalian 
species in Soviet Union. Two of the 3 suborders are represented (the 
third, Tylopoda, is present in a domesticated state), two of the 4 
infraorders, three of the 6 superfamilies, three of the 9 families, seven 
of the 15 subfamilies, and fourteen of the 86 genera. Artiodactyls 
thus constitute the most abundant group of ungulates in the Soviet 
Union. 

Artiodactyls are found throughout the Soviet Union but the 
number of species in the south is greater. The distribution of almost 



extinct and extant forms is, strictly speaking, not entirely logical since the very brief 
period of existence of extant forms in the evolution of groups is thus compared with 
the entire historic period. Precisely for the same reason placing extant and extinct 
forms together in a common taxonomic system should, at best, be tentative. 



all species underwent significant changes and shrank in the last 
century; some even became totally extinct or virtually so. In recent 
years the ranges of some species have been artificially expanded by 
acclimatization. 

Artiodactyls are found in the Soviet Union in all topographic 
zones and at altitudes exceeding 5,000 m above sea level. Most spe- 
cies are stenotopic (some strictly) and a few eurytopic. Some are 
typical gregarious forms while others live in small groups or even 
singly. In many species the herd instinct varies intensely throughout 
the course of a year. Some undertake significant seasonal migra- 
tions, while other species are stationary or shift only insignificantly 
in altitude. 

Nearly all these animals are distinctly polygamous. Mating usu- 
ally occurs in autumn and the young are born in spring. Some 
species possess a latent stage of development of the fertilized egg. 
Offspring generally number no more than two but sometimes reach 
10 or 12. Except for one, Soviet species are herbivorous. One or two 
molts occur each year. Seasonal variability is evident in several spe- 
cies, sexual dimorphism prominent in most, and age-related 
changes seen in some. 

In the Soviet Union artiodactyls play an important role in the 
hunting trade. They provide meat and hide and some yield fur and 
raw material for pharmaceutical and perfume industries and other 
consumer goods. Some products are exported and in some places 
play a vital role in the local economy. Some are maintained in a 
semidomesticated state; measures have been undertaken for the 
domestication of several. 

During the last century most species of artiodactyls underwent 
intense numerical reduction; some have since been placed under 
total or partial protection. In recent years the population of some 
species has risen and their range been enlarged as a result of such 
conservation measures. Artiodactyls represent an important group 
protected by conservation laws in most preserves. (V.H.) 

20 Key to Suborders and Families of Artiodactyla 

1 (4). Incisors present on upper jaw. 

2 (3). One pair of incisors present on upper jaw. Enlarged nails 

present in place of normal hooves; only two digits on legs 
(lateral ones absent). One or two fatty humps present on back. 

Domesticated animals 

Suborder TYLOPODA, Camel Family, Camelidae. 

3 (2). Three pairs of incisors on upper jaw. Normal hooves present; 



10 



four digits on legs (small lateral ones present). General 

appearance of swine 

Suborder SUIFORMES, Pig Family, Suidae. 

4 (1 ). Incisors absent on upper jaw 

Suborder RUMINANTIA. 

5 (6). Nostrils wide-set; distance between them more than shortest 

distance from lower tip of nostrils to edge of lip. Sometimes 
light-colored spots present on skin. Horns, if present, in the 
form of hard bony rods or covered with skin with short hair 
("velvet"), and usually branched (main beam with lateral 
tines). Teeth with relatively low crowns and vertical ridges 
and furrows on lateral surface do not reach edge of alveolus; 
ridges merge into horizontal ridge at base of crown (Fig. 2). 
Upper canines often present. Between frontal, nasal, lacrimal, 
and maxilla large, irregularly triangular vacuity present on 
each side Deer Family, Cervidae. 





Fig. 2. Left — Upper molar of deer (Cervus, family Cervidae); 

brachyodont with transverse ridge connecting outer ridges of crown 

at its base. Right — Same of goat (Capra, family Bovidae); hypso- 

dont without transverse ridge (from Sokolov, 1959). 

6 (5). Nostrils close-set; distance between their inner edges less than 
distance from lower end of nostril to lower edge of lip.^ Bright 
spots absent on skin. Horns, if present, in the form of horny 
sheath covering bony axis in the form of an out-growth of 
frontal bone and unbranched (main beam with no lateral 
tines). Teeth with relatively high crowns and vertical ridges 
and furrows on lateral surface reach edge of alveolus; horizon- 
tal ridge at base of crown absent. Upper canines absent. 
Between frontal, nasal, lacrimal, and maxilla vacuity absent 
or, if present, small and slitlike . . . Cattle Family, Bovidae, (V.H.) 



'Ratio differs in European wisents and domesticated yaks. 



Suborder SUIFORMES Jaeckel, 1911 



21 Artiodactyls are sometimes generalized but often specialized small-, 
medium-, or large-sized animals. In Suiformes the general build is 
massive and bulky, the limbs short, and the trunk massive and heavy. 

The number of digits is usually four; a few have three digits on 
the hind limbs. 

Reduction of the lateral digits is relatively poor; in some they 
differ morphologically and functionally only slightly from the 
medial and reach to the ground. All metatarsal and metacarpal 
bones fully developed and normally joined with tarsal and carpal 
regions. Fusion of metatarsal and metacarpal bones nonexistent or 
only rudimentary. Cannon bone absent. Terminal phalanges of dig- 
its covered with hooves. Ulna and (with many exceptions) fibula 
normally developed and free. 

Elongated facial part of skull usually wedge-shaped; in some, 
extended facial part greatly enlarged anteriorly. Longitudinal axis 
of skull more or less straight. Orbit opens posteriorly (with one 
exception). Ethmoid pit absent. Sagittal crest present. Horns absent. 
Mastoid bone covered. Articular condyle of lower jaw longitudinal. 

Canines invariably present and usually very well developed. 
Upper incisors usually number three and occasionally two or one. 
Molars bunodont or brachyodont, and exceptionally hypsodont. 

Stomach relatively simple and consists of not more than three 
sections. Suiformes are nonruminants. Cecum medium in size and 
simple or else absent. Sweat glands usually absent, present excep- 
tionally. Hair coat in some animals greatly reduced and bristly in 
most. Teats inguinal, or inguinal and abdominal, numbering one to 
six pairs. Placenta diffuse. Sexual dimorphism either not manifested 
or poorly so. Age-related changes generally absent. 

Suiformes are relatively slow-moving animals incapable of swift 
or long-distance running. Two major biological types are known: 
swine and hippopotami. They inhabit tropical to temperate forests, 
wooded zones of mountains, steppes, and marshy and fluvial 
regions. Most prefer extremely humid regions and one genus leads 
an amphibious life (Hippopotamus). They do not form significant 
herds and live in small groups, rarely singly. Offspring number one 

The pygmy hippopotamus {Choeropsis), in spite of common opinion, differs 
greatly in its mode of life from the common hippopotamus. 




3 w 



^ So 



13 

rarely, two occasionally, and generally ten to twelve. 
23 Most Suiformes are omnivorous with a predominantly herbivo- 
rous diet; some are purely herbivorous and feed mainly on aquatic 
vegetation. 

These nonruminants are widely distributed and occur in central 
and southern Europe and Asia, in the south up to the extreme points 
of habitat of the order, and in the north up to the southern part of 
the Scandinavian Peninsula and Gulf of Finland, Oka [River], 
southern Urals, southwestern Siberia, Altai [Mts.], Sayan [Mts.], 
northern extremity of [Lake] Baikal, and the lower Amur [River].* 
They are also found in North America in the north up to southwest- 
ern Texas, southeastern New Mexico, and southern Arizona. In Cen- 
tral and South America they occur in the south up to the equator, 
northern and northwestern Peru, Uruguay, and northern and cen- 
tral Argentina (up to the latitude of La Plata [River] and slightly 
southward). In Africa, they are found in northwest Africa, the Nile 
basin, and throughout the continent south of the Sahara, in Mada- 
gascar, and only in the northwestern Mediterranean Sea portion of 
the Arabian Peninsula (reconstructed range). 

Recent nonruminants differ distinctly from all other artiodactyls 
and there are no intermediate forms whatsoever between the various 
recent suborders. Morphologically, and partly biologically (omni- 
vorous feeding), the suborder of nonruminants, in spite of the high 
degree of specialization of some of its members, can be considered 
relatively poorly specialized and primitive. Forms at the commence- 
ment and in the first half of the Tertiary period point to a close 
phylogenetic relationship between nonruminants and the rest of the 
artiodactyl suborders. Keeping in view the infraorder Palaeodonta, 
some scientists regard nonruminants as the ancestral group giving 
rise to all other recent as well as extinct groups of artiodactyls. 

Members of the nonruminant suborder are already known from 
the Lower Eocene of America and Europe. They belong to the 
extinct infraorder Palaeodonta. Recent groups appeared not earlier 
than the Middle Eocene or Lower Oligocene (Europe). In the Ter- 
tiary period nonruminants developed relatively more abundantly 
than in the recent epoch. In the Eocene and Oligocene epochs they 
predominated, later giving rise to ruminants. At present they repre- 
sent a naturally declining group although there are some flourishing 
species among them. 



•The style of the Russian original often omits descriptive nouns from place- 
names; the reader should bear this in mind — Sci. Ed. 



14 

The suborder is divided into two* infraorders of which two are 
extinct (Palaeodonta and Oreodonta) and two extant (Suina and 
Ancodonta). Suina include one superfamily — Suoidea — with two 
families; Suidae — true hogs, and Tayassuidae — peccaries or New 
World hogs. The latter family consists of one (according to some, 
two) genus (Tayassu) with two species and is confined to America. 
These are unique nonruminants of the New World. There are no 
extinct families among Suina. 

Ancodonta include one extinct and one recent superfamily. The 
latter, Anthracotherioidea, consists of three families, of which only 
one is recent, i.e., Hippopotamidae. It consists of two monotypic 
genera — Hippopotamus and Choeropsis (pygmy hippopotamus) — 
distributed in equatorial Africa. 

The suborder consists of 134 genera (about 32%) of all artiodac- 
tyls) included in 8 families (about 32%). Of these, eight genera (about 
9.3% of recent artiodact'yls), included in three families (12%), are 
extant. 

The taxonomy of species of the suborder is simple. The total 
number of extant species is 12. Most (six) belong to Africa, two to 
America, three to Eurasia, and one is Eurasio-African. 

These are game animals providing meat, hide, and other mate- 
rial for making consumer goods ("ivory bone," hippopotamus 
teeth) and are essentially objects of sport. At some places they cause 
significant damage to plantations. One species is highly domesticated. 

Fauna of the Soviet Union include one of the two infraorders 
(Suina Gray, 1868), one of the two superfamilies (Suoidea Cope, 1887), 
and one of the three families (Suidae Gray, 1821). One species occurs 
in the Soviet Union (about 8% of the species of the suborder) repre- 
senting 0.3% of the mammalian species of this country. 

Suiformes are distributed in the southern half of the Soviet 
Union. 



*Misprint in Russian original; should read four — General Editor. 



Infraorder SUINA Gray, 1868 



24 Family SUIDAE Gray, 1821 (Swine) 

Suidae are nonruminants of medium size. 

All limbs with four digits each. Lateral digits well developed but 
distinctly shorter than medial ones, usually do not reach the ground, 
do not have a normal joint with the tarsal and carpal regions, and 
are markedly turned backward. Ulna and fibula free. 

As a result of elongation of mainly the maxillae and partly the 
premaxillae, skull elongated and wedge-shaped and not broadened 
anteriorly. Frontais, parietals, and supraoccipitals pneumatic. Orbit 
open. Facial part of lacrimal well developed and separates jugal 
from frontal or even frontal from maxilla. Usually two lacrimal 
foramina present. Hard palate elongated and extends backward 
beyond posterior edge of tooth row. Tympanic bulla swollen, pyri- 
form, and divided by bony septa into innumerable tiny chambers. 
Exoccipital and squamosal fused and mastoid set deep inside and 
not visible from the surface (covered). Dental formula (in the type): 
i — , с — , pm — , m -^ , but a reduction in number of incisors (to 
— ) and premolars (to -=-) occurs in different genera. Canines grow 
throughout life and upper ones turned upward. Incisors, premolars, 
and molars elongated posteriorly and typically bunodont and bra- 
chyodont; hypsodont molars and horizontal succession of teeth 
exceptional. 

Anterior end of snout extends into a small, relatively poorly 
mobile proboscis with a vertically slit anterior surface on which the 
nostrils open directly. Proboscis has a cartilaginous base which usu- 
ally ossifies (os nasi). Teats abdominal or inguinal and abdominal 
and number more than two pairs. Cecum present. Stomach simple, 
with only a small supplementary diverticulum ventriculi. Sexual 
dimorphism relatively poor. Age-related changes seen in some. 

Biologically Suidae are fairly monotypic, living in forests of dif- 
ferent types, in forest steppes and steppes, in the plains, and at 
moderate altitudes on mountains. They exhibit a preference for 
highly humid regions. Offspring number 4 to б and sometimes more 
(up to 10, and probably even 12). The animals are omnivorous with 
a predominantly herbivorous diet. 

25 The distribution of Suidae extends over part of the range of the 
suborder falling in the Old World. 



16 

Extant hogs represent a fairly monotypic, taxonomically well- 
defined group. They differ notably from hippopotami (different 
infraorder) but are highly similar to peccaries (family Tayassuidae). 
Sometimes both groups are merged into a common family with the 
rank of subfamilies. In reality the two groups are quite distinct 
(almost totally rudimentary fourth digit on the hind limb which is 
functionally tridactyl, fusion of proximal part of metapodia III and 
IV, special articulation of lower jaw, dorsal gland, etc. found in 
Tayassuidae). The two families are distinct even in the lower Ter- 
tiary. Peccaries are seen from the Lower Oligocene onwards and 
have developed independently in the New World where true hogs 
(Suidae) are absent and, in fact, were never present.^ Among non- 
ruminants true hogs represent an extremely primitive group, closest 
to the ancestral types of the infraorder Suina. Nevertheless, Tayas- 
suidae are characterized by several primitive features, even more 
primitive than those of Suidae. 

Representatives of these families, belonging to extinct subfami- 
lies, are known from the Lower Oligocene of Europe. The extant 
subfamily Suinae is seen from the Lower Pliocene onwards in 
Europe and Asia. 

Family Suidae is subdivided into five subfamilies, of which four 
are extinct (some survived up to the Pleistocene) and one extant, i.e., 
Suidae.* The family comprises 22 genera (about 16% of all nonrumi- 
nants) divided among five subfamilies (out of 19, i.e., 26%). Extant 
genera in the family number five out of eight in the suborder 
(62%): Sus — Eurasia, Potamochoerus, Hylochoerus, Phacochoerus — 
Africa, and Babyrousa — Sulawesi (Celebes). 

The total number of extant species in the family is eight, i.e., 66% 
of the extant species of the suborder or about 4% of the number of 
extant species of the order and about 0.3% of all extant mammals. 

Only one genus is known in the Soviet Union (20% of extant 
forms). (V.H.) 

I. BOAR, OR WILD SWINE 

Genus Sus Linnaeus, 1758 

1758. Sus Linnaeus. Syst. Naturae, 10th ed., vol. 1, p. 49. Sus scrofa 
Linnaeus. 



^Tayassuidae were present in Europe and Asia, continuing in Asia up to the 
Lower Pliocene. 

♦Error in Russian original; should read "Suinae"— General Editor. 



17 

Occipital region of skull highly raised and posterior profile ver- 
tical or slanted forward. Exostoses on skull and at base of upper 
canines absent or latter very poorly formed. 

Dental formula: i |, с \-, pm ^, m |-==44. Premolar ^ has no 
milk precursor and, like i - (even i y), tends to disappear (often absent 
in old animals). Medial incisors much larger than outer; lower incisors 
large and set almost horizontally. Canines grow throughout life; 
lower ones trifaceted. Canines distinctly smaller in females than in 
males. Tips of upper canines beveled due to grinding against lower 
ones. Molars brachyodont with relatively sharp cusps; succession of 
molars normal. 

Hair coat well developed, bristly in some and soft, furry in others; 
on neck and along spine hair long and sometimes very thick forming 
a prominent "mane". 

Teats six (Sus) or three (Porcula) pairs and abdominal and 
inguinal. 

Adults uniformly colored (sometimes with light-colored mark on 
snout). Young of most species with longitudinal light-colored 
patches on body. 

Habitats — forests, tugais [vegetation-covered bottomlands], and 
reed thickets. 

The geographic distribution of genus Sus (reconstructed) covers 
northwest Africa and Rio de Oro, Egyptian region of United Arab 
Republic, and Sudan (Sennar, Kordofan, and adjoining territories). 
26 In Eurasia the genus is found in the north in southern Sweden, Gulf 
of Finland, Oka, southern Urals, southwestern Siberia, and north- 
ern Baikal and Amur; in the south in Asia Minor, western Mediterra- 
nean, parts of Arabian Peninsula (Syrian region of United Arab 
Republic and Palestine), Ceylon, Nicobar and Andaman Islands, 
Sumatra, Java, Sumbawa, Sumba, Rota, Timor, Ara, Kai, New Gui- 
nea, Louisiade, Solomon Islands, Bismarck and Admiralty Islands, 
some small islands on the northern coast of New Guinea ( Yapen, for 
example), Waiges, Moluccas, and Philippine Islands. Hainan, Tai- 
wan, and the islands of Japan,^ including Ryukyu, also fall within 
the range oi Sus. 

^he natural geographic range of the genus in the extreme southeast is very 
difficult to demarcate, in fact, almost impossible. In the distant past local inhabitants 
of the islands, who were excellent sea-farers, imported domestic pigs (rather unique as 
domestic animals) which ran wild and now represent current wild forms. This is 
primarily true of New Guinea and the adjoining island groups occupied by S. 
papuensis. However, wild pigs exhibit signs of domestication in certain other areas of 
the islands between Australia and Asia, even for example Sulawesi (Celebes) (the 
entire group of celebensis type). Evidently in some places hybridization of truly wild 



18 




Fig. 4. Geographic distribution of extant wild boars, genus Sus L. 
(reconstructed). 

Gap occurs in range in central parts of Asia. Continuous line demarcates geographic 

range of common wild boar (Sus scrofa L.). Broken line separates that part of range 

where only Sus barbatus occurs. On some islands and in Malacca both Sus scrofa and 

Sus barbatus are found. V.G. Heptner. 



Genus Sus is distinctly separate from other genera of the family. 
It is closely related (through S. barbatus) to genus Potamochoerus 
(Africa, Madagascar). Genus Sus represents the least specialized 
group of recent hogs and is close to primitive types of the family; it 
is known from the Lower Pliocene of Europe and Asia. 

Genus Sus is very compact and contains so few species that 
attempts to divide it into several subgenera (Euhys, Aulacochoerus, 
and Dasychoerus) are unfounded. Only the separation of subgenus 
Porcula with one species (S. salvanius Hodgs.) appears justifiable. 

The taxonomy of species of genus Sus, unlike that of other gen- 
era, was extremely confusing until very recently. Seven, eight, and 
even more species with a large number of subspecies were acknowl- 
27 edged in the genus. Particular difficulties were posed by forms from 
South Asia, mainly the island region between Asia and Australia. 

indigenous local hogs with the imported forms which ran wild occurred. It is highly 
possible that the true geographic range did not extend so far southeast as shown here. 



19 

Apart from other reasons (incomplete descriptions and the like), 
complexity of development of structures in these forms made classifi- 
cation difficult. In addition to phenomena associated with island 
life, a powerful influence has been exerted by hybridization of 
domestic and wild forms and the transformation of the former into 
the latter. 

In recent years the situation has been clarified somewhat and 
only three species should properly be placed in genus Sus: S. scrofa 
(including the complex of S. leucomystax from east Asia, S. cristatus 
from India and Indochina, S. vittatus from Sumatra and Malacca, 
and some others), S. barbatus (including verrucosus, celebensis, and 
some others) from Malacca, Sumatra, Java, Kalimantan (Borneo), 
Sulawesi (Celebes), Moluccas and Philippine Islands, and S. salva- 
nius, the pygmy hog from Sikkim, Nepal and Bhutan. 

Genus Sus thus includes about 25% of the species of the family. 

Only one species is known in the Soviet Union (33% of extant 
species of the genus), representing 0.3% of the mammalian species of 
the Soviet Union. (V.H.) 

1. Sus (Sus) scrofa Linnaeus, 1758 

1758. Sus scrofa. Linnaeus. Syst. Naturae, 10th ed., vol. 1, p. 49. 
Germany. 1811. Sus europaeus. Pallas. Zoographia Rosso-Asiatica, 
vol. 1, p. 265. Caucasus to Baikal, substitute for Sus scrofa Linn. 
1875. Sus scrofa var. nigripes. Blandford. Journ. As. Soc. Bengal, 
vol. 44, pt. 2, p. 112. Tien-Shan in Kashgar. 1888. Sus ussuricus. 
Heude. Mem. cone. I'Hist. Nat. Emp. Chin., vol. 2, p. 54. Ussuri 
valley. 1889. Sus leucomystax var. continentalis. Nehring. Sit- 
zungsber. Ges. naturf. Freunde, Berlin, p. 141. Vladivostok. 1892. 
Sus gigas. Heude. Mem. cone. I'Hist. Nat. Emp. Chin., vol. 3, p. 191. 
Vladivostok. 1912. Sus attila. Thomas. Proc. Zool. Soc. London, p. 
183. Kolozvar (Klausenburg, now Cluj) in Transylvania (Rumania). 
1918. Sus falz-feini. Matschie. Sitzungsber. Ges. naturf. Freunde, Ber- 

''This species (height at shoulders about 30 cm and skull length 150 to 160 mm), 
described as early as 1847, was long regarded as non-existent and merely a name used 
to describe young or abnormal or underdeveloped animals. The independent exist- 
ence of this species was demonstrated only recently (Ellerman and Morrison-Scott, 
1951). 

The enigmatic form S. gargantua, a gigantic hog of Borneo known only from a 
single skull, is not included here. 

^Young animals, up to one year of age, irrespective of sex termed "sucklings", and 
from one to two years "juveniles". Adult males called "boars". Fully grown adults 
with good canines (tusks) called "tuskers" and old males with tusks "loners". Adult 
females called "sows". 



20 

lin, no. 8, p. 1. Naliboki, about 75 km west of Minsk. 1922. Sus 
leucomystax sibiricus. Staffe. Arb. d. Lehrkanz. Tierzucht. a.d. 
Hochschule f. Bodenkultur, Wien, vol. 1, pp. 51-58, Pis. I-III. Tun- 
kin mountain range, southern Siberia. 1930. Sus scrofa raddeanus. 
Adlerberg. Doklady Akad. Nauk USSR, p. 95. Sugu-nor, southern 
Hentei, Mongolia. (V.H.) 

Diagnosis 

Sus {Sus) scrofa is the only species of genus Sus found in the Soviet 
Union. 

Description 

General build massive and bulky with short and relatively thin legs. 
28 Trunk relatively short with an extremely massive anterior part and 
poorly developed posterior part. Back in region of shoulder blades 
highly raised (humped) and neck thick, short, and nearly immobile. 
Head very large and constitutes about one-third body length (includ- 
ing head); projects forward like a wedge and continues inconspicu- 
ously into neck and withers. Tail thin and short, not reaching hock, 
with a few elongated hairs at tip. 

Ears long and broad. Eyes small and deep-set. Snout projects; 
nostrils set directly forward, open on broadened, blunt edge of snout. 
Upper and lower canines protrude from mouth of adult males. 

Hooves of medial digits larger and elongated, with pointed tips 
and edges bent downward, capable of quick movement; hooves of 
lateral digits also well developed, set low, and relatively large, long, 
and sharp.^ 

In general appearance females somewhat lighter in build and 
more slender than males, with a smaller hump and less massive 
anterior trunk portion. 

In winter wild boar are covered with long, dense, resilient, some- 
what curly bristles underlaid with an extremely short, dense lining 
of fur consisting of thin, curly, soft, downy, light brown or brown- 
colored hair. The base of each bristle is dark-colored (black or 
blackish-brown) and the terminal part lighter. The bristles on the 
head are short, longer on the sides, especially on the back, and long- 

*Compared to other Russian ungulates the lateral digits and their skeleton are less 
reduced and more functional in wild boar. The metapodium represents a compact, 
greatly perfected supporting system, especially when moving on soft ground. When 
resting on all four digits, the weight load on 1 cm^ is roughly reduced by 1.5 times, 
i.e., from 500 to 320 to 390 g (Sludskii, 1956). 



21 




Fig. 5. Tusker. Sketch by A.N. Komarov. 



est in the middle of the -back in the anterior part of the body. A 
29 characteristic crest forms here and on excitation these bristles stand 
erect. Both the abdomen and legs are covered with very short bristles. 
The latter are directed forward on the rear of the abdomen and in the 
lower part of the neck, and directed backward on the rest of the body. 
The overall color of the animal depends on the hue of the hair 
(bristle) tips and is highly variable. Wild boars in Balkhash vary 
from light brown to pale yellow (sandy) or even w^hite. In Belorussia 
rusty-brown-colored animals, lighter-colored animals, and even 
totally black ones are seen. In Ussuri region ocherous, light brown, 
and black individuals are found. Extremely light-colored animals 
have the silver-gray color of badgers. In a new coat individual body 
parts vary in color as follows: posterior part of head, neck, back, 



22 




Fig. 6. Sow with sucklings. Sketch by A.N. Komarov. 



sides, and tail relatively light and anterior parts like the snout and 
ears, in addition to the legs dark, sometimes black. In some forms 
(see below) a light-colored patch runs from the corner of the mouth 
backward. Hooves are always black. 

By spring the hair cover, especially in central Asia and Kazakh- 
stan, has usually undergone intense fading and the color particu- 
larly of sucklings and juveniles is notably lighter. In summer, when 
the winter hair is shed and the new coat has just begun to grow (one 
molt occurs in wild boar, see below) and the fur lining has not yet 



23 

formed, wild boar appear very dark, often dark brown and black, or 
black. This is mainly because the black skin shows through the 
sparse hair coat. Hooves and snout are black. There are no sex- 
related variations in color. 
30 Color varies sharply with age, however. Sucklings retain the rela- 
tively soft juvenile hair with which they are born and their general 
color is light brown or rusty-brown. Longitudinal, light-colored, 
pale yellow bands extend along the sides and back. The adult coat 
develops in the fourth month after birth. Juveniles, at least in some 
areas, are usually more lightly colored than adults and older 
individuals. 

Some geographic variability is also evident, in addition to dis- 
tinct individual variations of pigmentation (see below). 

The skull of the male wild boar is massive and characterized by 
great development of upper and lower canines which grow through- 
out life. The upper ones are massive with a blunt apex and edges, 
relatively short, triangular in cross section, initially grow sideways 
and later curve upward; in older animals they are turned backward. 
The lower canines are also triangular in cross section but much 
longer (their exposed part is 10 to 12 cm or more long ), turned back 
like a saber, with sharp cutting edges and a very sharp apex. A 
prominent growth along the edge of the jawbone occurs in the basal 
region of the upper canines. 

Canines begin to grow in the male even in the first year of life 
and are well developed by the age of three years; they continue to 
grow throughout life but are somewhat flexed backward in later 
years, especially the upper ones. Canines are poorly developed in the 
female and do not serve as tools; they are of little importance in 
burrowing, rooting, and so on. 

Sequential changes linked with age are distinct in the skull with 
advancing age; the relatively short facial region of the young 
becomes greatly elongated. 

In August, before the breeding period, a "stuffing" (also called 
"fat" or "armor") of subcutaneous tissue resembling cartilage forms 
along the sides of the body of the male and protects him during 
fights. This tissue covers the shoulder blade region and extends 
backward to the last rib; it is thickest (2 or 3 cm) in the region of the 
shoulder blades and gradually thins backward toward the rump and 
forward toward the neck. Dorsal to the preputial orifice a special sac 

Some record specimens of canines are much longer — 24 or 25 cm along the outer 
curvature including the portion embedded in the bone. A large lower canine has a 
nearly regular semicircular form when removed from the socket. 



24 

the size of a large egg when extended or even sHghtly larger (diverti- 
culum preputiale) also occurs in the male. Some urine collects and 
settles in it giving off a sharp, unpleasant odor. The function of this 
diverticulum and its characteristic musculature is not entirely 
understood; it evidently has a mechanical function and helps pre- 
vent the outflow of blood from the corpus cavernosum.* Hesse's 
cardiac index is very low — 4.4 to 6.5 (Bannikov, 1955). 

Wild boar vary considerably in size in accordance with age. They 
grow slowly, attaining full growth at the age of five or six years; 
even then growth does not cease. Sex-related differences are also 
notable; females are invariably smaller than males. Weight also 
changes, depending largely on the degree of feeding during different 
years and seasons. During the breeding season males become highly 
emaciated. Size and weight also reflect some geographic variability. 

The size of adult boars (not younger than five to six years) from 
the lower reaches of the Hi (Semirech'e) is as follows (in cm)^: body 
length: males 148 to 174 (M* 160) (n=ll), females 127 to 157 (M 144) 
(n=12); length of torso: males 90 to 118 (M 98) (n = 11), females 80 to 
108 (M 90) (n = 9); height at withers: males 79 to 102 (M 86) (n = 11), 
females 80 to 86 (M 77) (n = 12); height at rump: males 78 to 102 (M 
31 80) (n = 11), females 70 to 81 (M 74) (n = 10); tail length: males 20 to 
29 (M 24.5) (n = 11), females 19 to 26 (M 22) (n = 10); length of ears: 
males 13 to 16 (M 14) (n = 11), females 12 to 14 (M 13) (n= 11); chest 
circumference: male 113 (n = 1) and female 95 (n = 1) (Sludskii, 1956). 
The average body length of western (European) wild boar is about 
150 cm and height at withers about a meter. 

The weight of Latvian wild boar may reach 200 or even 236 kg 
(Kalnin'sh, 1950) and of Belorussian animals 256 kg, although boar 
usually weigh about 150 kg or slightly more (Serzhanin, 1955). Car- 
pathian animals weigh 120 to 180 kg and exceptionally up to 250 
(Tatarinov, 1956). Wild boar in the Caucasian preserve in the 1930's 
weighed as follows: males 64 to 178 kg (M = 166 kg) and females 48 to 



'in any case it is not a musk gland as reported by some authors (Sludskii, 1956). 

'The measurements given for Semirech'e wild boar are far from maximum values. 
Evidently they often reach as much as 200 cm in length. At the same time references to 
body lengths of 230 and 240 cm and height at withers of 1 10 cm or more, especially of 
western animals (Latvia and Belorussia; Serzhanin, 1956; Kalnin'sh, 1950) are dub- 
ious or relate only to exceptional specimens. 

*Here and elsewhere "M" stands for mean measurements — General Editor. 

'^Footnote indicated but none given in Russian original— General Editor. 



25 

108 kg (М = 68 kg) (Donaurov and Teplov, 1938); early in this 
century old males weighed 240 to 255 kg and females 120 to 145 kg 
(Dinnik, 1919). Animals of the Astrakhan' preserve (Volga delta) in 
the 1 930 's weighed up to 250 to 260 kg (males; Lavrovskii, 1952) and 
wild boar in Syr-Darya up to 240 kg. A few years earlier animals of 
270 kg and even 320 kg were reportedly caught. Wild boar of Hi 
touch the scale at 220 to 240 kg and sows at 95 to 100 kg (Sludskii, 
1955). In Eastern Siberia wild boar attain weights of 240 or even 272 
kg. Boars of the Far East are particularly heavy; old tuskers tip the 
scales at 300 to 320 kg (Abramov, 1956). 

As can be seen from the data available, the Caucasus wild boar 
used to be larger than they are at present. The weight of animals 
from the northern Caspian coast in the middle of the last century 
went up to 320 kg. Reduction in size of wild boar has been reported 
nearly everywhere in the world; 200 pood* animals are now rare 
even in the Far East where boar weighing 200 kg are now considered 
large. This is the result of thoughtless hunting, due to which the 
average age of the animals is lowered since they are not allowed to 
grow to maximum size. It is significant that in the Astrakhan' pre- 
serve in the 1940's the animals were notably larger than at the end of 
the last century when they were numerically very abundant. At that 
time they attained a maximum weight of only 192 kg versus 250 to 
270 kg at present (Sludskii, 1956)." 

In Semirech'e the live weight of sucklings at the age of 8 to 11 
months is 21 to 30 kg for males and 20 to 29 kg for females, i.e., at the 
time of completing their first year suckling weight is about one- 
fourth the adult weight. Their body length at that time is 93 to 115 
cm in males and 92 to 105 cm in females. Young pig between the 
first and second year of life weigh 25 to 54 kg and 35 to 44 kg (male 
and female respectively); this represents roughly one-half or slightly 
less than the average weight of adult animals of corresponding 
sexes. The body length is 120 to 130 cm and 100 to 120 cm (male and 
female respectively) (Sludskii, 1956). Newborns weigh 1,620 and 
1,360 g respectively (Ussuri region; G.F. Bromlei). (V.H.) 

Taxonomy 

Within the genus S. scrofa is closest to bearded swine (S. barbatus 
Mull.) inhabiting Malacca and other islands (see Fig. 4 and descrip- 

*Pood= 16.38 kg— General Editor. 

"Mean weight of male from Central Europe 130 kg; maximum evidently about 
230 kg. 



26 

tion of geographic range of wild boar). The third species of the 
genus — 5. salvanius — found in a very narrow region in the Hima- 
layas and possibly a relict form, stands quite isolated and is usually 
placed in a separate subgenus or sometimes even a separate genus, 
Porcula. (V.H.) 

Geographic Distribution 

The geographic distribution extends in Europe up to 60 to 55° N lat; 
in Asia up to 55 to 50° N lat. and in the south includes India and 
Ceylon and a significant part of the island zone from Malacca to 
Solomon Islands, and northeastern and northwestern Africa. 

Geographic Range in the Soviet Union 

The range in the Soviet Union covers the extreme northern section 
of the range of the species. It extends over the southern half of the 
European part of the country, the Caucasus, and the southern 
fringes of the Asiatic part of the USSR. 
32 The range of wild boar in the Soviet Union has varied signifi- 
cantly over the years. This can be seen from the fact that almost the 
entire northern fringe of the range falls within the Soviet Union and 
that the species has survived in very large expanses under minimal 
or close to minimal climatic (temperature and snow) and trophic 
conditions. Under such conditions the lability of the fringe of the 
range and its boundaries is readily apparent. The human factor has 
also played a major role in changing the range of modern boar. 
Foremost among the various causes are hunting the animals and 
destruction of their biotopes. Against the background of minimal 
conditions such factors become acute. It is very easy to eliminate 
wild boar populations and very difficult to restore them. 

In the last century the range of wild boar shrank extensively in 
the European part of the USSR. The animal disappeared almost 
throughout the territory except in the far west and in a negligible 
section in the southeast (see below). In the region from the Ural 
River to the Altai also the boundary has fluctuated extensively and 
shifted far to the south. A particularly intense destruction of animals 
and reduction of range occurred in the second half of the nineteenth 
century and in the first few decades of the twentieth. The range was 
reduced to its minimum evidently in the 1930's. 

'^It is possible that the process was somewhat different in the Baltic region and the 
second half of the nineteenth and the beginning of the twentieth centuries represent a 
period of expansion of the range in the north (Ling, 1955). 



27 

A noticeable expansion of the range of wild boar commenced in 
the 1930's in the European part of the Soviet Union and western 
Altai. Expansion proceeded very rapidly between 1940 and I960, 
expecially in Kazakhstan. However, in other regions, for example 
northern Caucasus, range expansion toward the north was not seen. 
By the mid-1950's the range boundary had advanced significantly 
but extended no farther since adequate numbers could not be formed 
in the newly occupied territories. In many places permanent habita- 
tion is still not seen and wild boar is an intruder. The process of 
colonization of new territories is still going on. Further expansion 
depends on climatic conditions in future years and, most important, 
on human activity. It is significant that under the present conditions 
of establishment of wild boar the boundaries have reached their 
former maximum limits in the north in several places. 

Reconstructed geographic range. The reconstructed northern 
boundary of the range of wild boar commences in the west at the 
eastern tip of the Gulf of Finland and southeast coast of Lake Lad- 
oga (Mga region).'^ This is the northernmost point of habitation of 
wild boar in the USSR (60° N lat.). From there the boundary steeply 
turns south to Novgorod and farther into the region of Lake Peno 
and the town of Ostashkov to the environs of Kalinin. From there 
the boundary sharply descends south running through the western 
fringes of Moscow district or even Moscow (intrusions). Finally, the 
boundary proceeds toward Kaluga and turns to Tula, running later 
to the east, probably south of Ryazan'. 

It is highly possible that wild boar occupied forest sections south 
of the Oka ("Trans-Oka Stockade"). From Ryazan' ("Ryazan' prin- 
cipality") the boundary of the range inexplicably turns east through 
Penza district ("Penza province," 18th century) and runs toward the 
Volga at Samarsk Luka or, probably, even more northward (wild 
boar is indicated in "Simbirsk province"). Wild boar reached its 
northern limit on the Volga probably by penetrating from the south, 
evidently along valleys of rivers like the Don and its tributaries 
(Khoper). At places far removed from the rivers the boundary could 
lie more to the south. 

East of the Volga the northern boundary of the range of wild 
boar runs from Zhigula (or Ulyanov district) toward the Buzuluk 
River, southern Ural, Ik, and Sakmar, then to the upper reaches of 



'^The present discussion pertains to wanderers and not permanent habitation. 
The separation of the region of wanderers from that of regular habitation is very 
difficuk. The references here pertain to peripheral points where^the animal is found 
irrespective of the nature of its habitation unless specifically stated otherwise. 



29 

the Beloe (Voznesensk Zavod — Irgizla; Kirikov, 1935). In the south- 
ern Urals wild boar reached in the north up to 52° or almost 53° N 
lat.''' After crossing the southern extremity and foothills of the 
Urals, the boundary in Trans-Urals runs toward Bashkir preserve 
(west of Magnitogorsk) and Magnitogorsk and rises northeast 
toward 55° N lat. There wild boar has been recorded on the line 
between Miass (west of Chelyabinsk) and Kurtamysh (slightly south 
of Kurgan) at 55° N lat. or slightly more northward. More eastward 
the northern point of habitation of wild boar lies at Mangut railway 
station at 55°45'N lat., southeast of Ishim town and even slightly 
more north to the east to Ishim town (Lakes Ik and Kuban, a little 
north of 56° N lat.). Farther away, the boundary crosses the Irtysh 
and Ust'-Tara at 56°33'N lat. and runs between the Irtysh and Ob' 
(Barabin steppe) covering Lake Chany and probably its basin, i.e., 
along 55° N lat. or slightly more north almost along 56° N lat. 

From the eastern part of Barabin steppe (Novosibirsk region) the 
boundary of the range turns sharply south into the region of the 
confluence of Biya and Katun, i.e., it almost approaches the Altai 
range. From there the boundary line encircles the Altai, turns 
south-east, passes into Zmeinogor region, and southward passes 
through the basin of the Tigerek River and, evidently, also Tigerek 
mountains or a part of them. Farther away, joining the Altai from 
the south, it proceeds southeast and east through the basins of the 
Shul'ba, Ul'ba, and Bukhtarma, and extends into China. '^ 

Small sections of the southern Altai-Chuisk steppe (upper 
reaches of the Chuya with the center in Kosh-Agach) and the 
Kuraisk steppe (below, along Chuya in the region of 87°40' to 88° E 
long.) and the upper reaches of Bashkaus (Chulyshman tributary) 
also fall in the range. Throughout this region wild boar was seen 



'''it is possible that the boundary beyond the Volga traverses somewhat even more 
northward than pointed out above but this is not supported by reliable data. Informa- 
tion about habitation up to Kama and Elabuga in Kama (Sludskii, 1956) is not 
sufficiently accurate and pertains to a long-past period (the ancient Anan'insk town- 
ship). The line of reconstructed northern boundary of the range in the European part 
of the USSR differs greatly from that proposed recently (Sludskii, 1956) and ran very 
far north from Ladoga through Rybinsk reservoir, north of Kostroma, through mid- 
dle Unzha, middle Vetluga, lower Vetluga, and Beloe estuary. The relative absence of 
wild boar at Vetluga even 1,(Ю0 to 2,(Ю0 years ago is a direct inference (Formozov, 
1946). 

''wild boar were also encountered evidently along the mountain slopes surround- 
ing the valley from the north (Muyuta River; Kolosov, 1939) but the assumption of 
their habitation on northern slopes of Katun range (Sludskii, 1956) can hardly be 
substantiated. 



30 

evidently only periodically as transgressions from Mongolia.'^ 
Farther away in the east the range covers all of Tannu-Ola up to its 
eastern extremity (Sangilen range) and the montane parts of Tuva in 
the upper parts of Little (Ka-Khem) and Great (Bii-Khem) Yenisei in 
the north up to the tributaries of the Great Yenisei, Azas, and Kham- 
sara (the latter runs from the southern slopes of Udin range to the 
point of its confluence with Okin), adjoining Pri-Kosogol district of 
Mongolia. The Tuvin section of the habitat in the south as well as 
in the east is associated with the range of wild boar in Mongolia and 
evidently forms a common unit with it. Within the Soviet Union it 
is isolated by high mountains or only weakly linked with the section 
of the habitat of the animal more to the north. 

The habitat of wild boar on the northern slopes of the eastern 
Sayan and to the north of this commences in the west, in the basin of 
the upper reaches of the Kungus tributary of the Kan (about 95°40' 
to 96° E long.). Its southern boundary runs along the northern foot- 
hills and the slopes of the above mountain system through the upper 
reaches of the Tagul and Biryusa, upper reaches of the system the 
Uda, li and Oka, Bol'shaya and Malaya Beloe, Kitoi and Irkut, 
encompassing the Tunkin range and passing along Irkut right up to 
Lake Baikal. This is indeed the southern boundary at present. No 
data are available about its former location but probably it differed 
little from that described above. There is very little information 
35 about the northern boundary of the former habitat at Baikal. It may 
be assumed to have extended to the Angara around 57° N lat. (edge 
of Dolgii region) from a more western section than indicated, possi- 
bly through Taishet region. Turning from there toward Baikal the 
boundary evidently inclined considerably south but covered the 
upper reaches of the Lena in the north up to Ust'-Ilga and even 
Ust'-Kut (56°30'N lat.; transgressions). Farther away, the Baikal 
shore right up to its southwestern extremity formed the boundary. 
In southern Baikal and beyond the boundary is now formed by 
the southern and eastern shores of the lake. From its northern 
extremity (probably from the upper reaches of Barguzin) the bound- 
ary proceeds north reaching Bodaybo region (northernmost point 
of the habitat in Siberia, around 57° to 57°30'N lat.). From there, 
the boundary inclines slightly south and proceeds east into the 
source of the Chara (57° N lat.). In the east it runs in the upper 
reaches of the Nyukzha River, a tributary of the Olekma, and pro- 

'*As a rare exception wild boar might possibly occur even farther north, deep 
inside the Altai taiga, since the skull of a wild boar was found in LakeTelets (Kamgin 
Bay; Zalesskii, 1934). 



31 

ceeds along 54° 45' N lat. to the Ol'doe (Amur tributary). From there 
the boundary descends somewhat south and encircles the Tukurin- 
gra range from the south, passing along its slopes and intersecting 
the left tributaries of the Urkan River, a tributary of the Zeya, at 
54°20', 54°30', and 54° N lat., and leaving the Zeya slighdy south of 
54° N lat. Farther east the boundary descends even more southward, 
intersects the upper reaches of the Bureya, encompasses the lower 
and middle reaches of the Urma and Kur', and turning parallel to 
the course of the Amur runs to the Gorin River slightly south of 
Lake Evoron. Beyond the Gorin the range extends along the Amur 
up to 51°30'N lat. (up to the estuary of Khazelakh River), encom- 
passes the region of the rivers Yai and Kod and Lake Kiza, and runs 
toward the Tatarskiy Strait at De-Kastri or slightly more south of it 
(Schrenk, 1859). It is possible that in the region south of the Gorin 
wild boar was an irregular visitor or an intruder.''' 

At Sakhalin wild boar is absent and was probably never present 
even in the historic past. Fossil wild boar have reportedly been 
found there, however. 

The natural range of wild boar in the Soviet Union is essentially 
a single continuous area, the gaps in the Altai-Sayan mountains 
being altogether insignificant; connecting points occur in the south 
in the immediate proximity of the present boundaries.'^ 

South of the above northern boundary the range extends every- 
where up to state boundaries and up to the coasts of the Azov and 
Black Seas and Crimea (in montane Crimea wild boar disappeared 
only in the middle of the nineteenth century) and even beyond them. 
The high montane areas of the Great Caucasus, Trans-Caucasus, 
Pamir, Pamir-Alai and Tien Shan, some mountains of Siberia, and 



'^According to other data for the middle of the last century (Maak, 1859), the range 
extended beyond the Gorin for an extremely short distance; the boundary on intersect- 
ing this river soon turned southeast and ran onto the right bank of the Amur slightly 
below the Gorin estuary. Running in the same direction, it extended toward Tatars- 
kiy Strait south of Kiza. 

'^The boundary of the reconstructed range has been ascertained from Kalnin'sh, 
1950; Ling, 1955; Nasimovich, 1955; Formozov, 1947; Satunin, 1895; Barabash- 
Nikiforov, 1956; Ptushenko, 1937; N.A. Severtsov, 1950; Kirikov, 1952 and 1955; 
Rychkov, 1762; Pallas, 1786; Eversmann, 1850; Gebler, 1837; Zalesskii, 1934; Makarov, 
1949; Kolosov, 1938 and 1939; Yanushevich, 1952; Favorskii, 1936; Podarevskii, 1936; 
Kopylov, 1948 and 1950; V.N. Skalon, Shcherbakov, and Bazykin, 1941; V.N. Skalon, 
1951; Maak, 1859; Rul'e, 1845; Middendorff, 1869; Radde, 1862; Gassovskii, 1927; 
Bannikov, 1954; Schrenk, 1859; Abramov, 1954; Serzhanin, 1955; and other sources; 
and according to material compiled by Nasimovich, 1955; Formozov, 1946; and Slud- 
skii, 1956. 



32 

Sikhote-Alin form gaps in the range. The central and deep interior 
parts of the desert (Karakum, Kyzylkum, and others) do not fall 
within the range although in winter, when moisture is abundant, 
wild boar in some places colonize areas deep (up to 100 to 200 km) 
inside the desert (V.G. Heptner; Sludskii, 1956). 

Wild boar belongs to the group of ecologically versatile species. 
As a common animal it is capable of withstanding very low temper- 
atures and surviving in areas with mean January temperatures of up 
to 20 to 25° С and even 30° С [below zero].* At the same time, because 
of its short legs and consequently its ability to utilize only food lying 
on the ground (or in it), its distribution is associated with some 
definite conditions of snow cover; it cannot normally exist in areas 
with a very high snow cover. '^ A snow height of 30 to 40 cm and not 
more than 50 cm, appears to be the limit. 
36 In the European part of the USSR the range boundary at some 
places coincides very precisely with the isoline of 50 cm snow depth. 
This line determines the boundary in the northwest between Rya- 
zan' district and the Volga and from the Volga to southern Urals. 
The significant diversion of the range boundary south in the area 
between the Volga and Oka Rivers and its position south of the Oka 
are due to excessive snow in this region. In Siberia, due to the 
extreme severity of environmental conditions, such correlations are 
more complex and less distinct and the picture in some respects less 
clear than in European USSR. However, the distinct curvature 
in the boundary to the north in the Baikal and Trans-Baikal regions 
coincides with the poor snow in these regions, while the descent of 
the boundary to the south in the Amur and Ussuri regions, on the 
contrary, correlates with the occurrence of heavy snow and so on. 

In addition to the association of wild boar with snow, the influ- 
ence of other factors on the range is also significant (see the section 
"Biology"). In a geographic sense it is particularly significant that 
the animal succumbs to destruction by man most easily in regions 
with abundant snow and mostly in years of particularly heavy snow- 
fall. All this occurs mainly in the northern, primarily in the border- 
line regions of the habitat of the animal, which in many cases could 
explain the boundary fluctuations of wild boar and its general 
recession. 

Geographic range in 1930. The range described above, as pointed 

♦Unspecified in Russian original — Sci. Ed. 
For information on tfie rorrelation of the range with snow see Middendorff, 
1869; Podarevskii, 1936; Formozov, 1946; Abramov, 1954; Nasimovich, 1956; Sludskii, 
1956; and others. 



33 

out, underwent intense shrinkage during the last century. The rea- 
son for this was destruction of the animal through thoughtless hunt- 
ing. Natural causes, i.e., climatic fluctuations (considering the 
phenomenon as a whole), per se played a minor role or acted only as 
factors promoting destruction. 

The recession of the northern boundary and the reduction of the 
range proceeded fairly rapidly, but became particularly intense from 
the middle of the last century. The process reached its culmination 
during the first one-third of this century. Boundary recession was 
least in the extreme northwest and Far East section of the range and 
comparatively insignificant in eastern Siberia. Recession was 
intense in western Siberia and Kazakhstan and significant changes 
occurred in European USSR (apart from the extreme western 
regions). 

By the beginning of the 1930's, which could be considered the 
period of maximum reduction of the range, the area not only 
decreased notably, but that which was formerly a continuous zone 
disintegrated into a few individual discontinuous or even isolated 
portions. This is true of all European USSR and the Asiatic region 
in the east up to the Altai. Changes in the east were less significant 
or altogether absent. 

The former zone of habitation of wild boar, as seen in 1930, 
covered far western parts of European USSR. Its northeastern bound- 
ary commenced roughly at Riga, turned toward Velikie Luki, and 
shortly before entering them turned south into Vitebsk or bypassed 
it from the west. Later, it ran east of Mogilev through Gomel' or 
somewhere between Gomel' and Bryansk (close to the former) and 
after covering the Chernigov region turned west, bypassing Kiev 
from the north. In a manner not precisely known it later turned 
southwest toward the upper Dnestr roughly at Mogilev-Podol. On 
its way to the Dnestr the boundary ran along the Black Sea and 
crossed westward along the USSR border into the estuary of the river 
along the sea coast (Bobrinskii, 1944; Formozov, 1946; Sludskii, 
1956). 

East of the above line demarcating the region, which represents a 
fairly permanent habitat of the animal, some rare encounters, some- 
times at quite distant places, have been recorded. Thus wild boar 
have been sighted in Ostashkov region. Lake Peno, Smolensk, 
Bryansk, and even Moscow district. 

The second habitat of wild boar covered the Caucasus and 
northwestern and northern banks (partly) of the Caspian Sea. The 
northern boundary of this part of the range ran from the Sea of Azov 
along Lake Kubana to Krasnodar, from there to Armavir, turning 



34 

around Mineral'nye Vody from the south, toward Prokhladnyi 
(slightly northeast of Nal'chik). From there, along the Terek, the 
boundary ran right up to the sea. From the Terek estuary, in a strip 
along the sea coast, the range extended north into the Volga delta 
and farther east up to the Ural estuary. This zone was very narrow 
and bounded nearly exclusively by a zone of Caspian reeds. Only at 
some places, for example, in the lower reaches of the Kuma, did it 
enlarge giving rise to small pockets away from the sea. Moreover, 
this band of the range was interrupted at some places as, for exam- 
ple, in the stretch between the estuaries of the Kuma and Volga. 

Beyond the Caspian Sea the range covered the Kopet-Dag, Great 
Balkhan, and the course of the Atrek except for its lower dried up 
part (roughly up to Lake Delili), the basins of Tedzhen and Murgab, 
and the extreme south of the interfluve of these rivers. Farther away 
the boundary covered the Amu-Darya from the state boundary to its 
left bank up to the Aral Sea and ran along the southeastern and 
eastern coasts of the Aral Sea up to the estuary of the Syr-Darya. 
From this river the boundary ran into the lower reaches of Lakes 
Sarys and Telekol'sk, encompassed the Chuya, ran along [Lake] 
Balkhash, and covering the Alakol', Tarbagatai, and the Zaisan 
depression, continued toward China. 

Small isolated sections of fairly permanent habitations of wild 
boar have been preserved north of the above line in Kazakhstan. 
These sections were isolated from the main range and comprise the 
area of Lake Chelkar-Tengiz and the lower reaches of the Turgai 
and Irgiz, Lake Kurgal'dzhin and Tengiz southwest of Akmolin, and 
perhaps, Kamysh-Samarsk lakes and the lower reaches of the Uze- 
nei-'^ (Bobrinskii, 1944; Sludskii, 1956; V.G. Heptner). 

Wild boar occurred in the southern Altai in the period under 
discussion only as an intruder in some of the areas mentioned above 
adjoining Mongolia. 

In the Sayan, west of Baikal, at the commencement of the 1930's 
the northern boundary ran much southward than it does now, and 
the range in general represented a fairly narrow strip along the 
Sayan foothills. Thus in the region of Biryusa it fell far short of 
reaching Taishet and the Uda basin to the north and did not cross 
54°33' N lat. From the li, in the basin of which wild boar proceeded 
north roughly up to 54° 10' to 54°20' N lat., the northern boundary 
turned southeast toward Baikal exiting on its southwestern extrem- 



^"in the region of the Ermentau highland wild boar was seen in the period under 
description only as an intruder from Kurgal'dzhin (Sludskii, 1956; map; pp. 168 and 
185). Pockets of habitation there (ibid., map; p. 147) were evidently absent. 



35 

ity or slightly closer to the Angara estuary. However, swine intru- 
sions have been reported at Angara (Kureika River, 54° 45' N lat.) 
and even at the sources of the Lena. 

In trans-Baikal at this time the range boundary receded south- 
ward and did not extend north of Kalar 56° N lat.) where the animal 
was extremely rare. Also, it did not cover the upper Angara (Poda- 
revskii, 1936; Kopylov, 1950; Troitskii, 1930; Skalon, Shcherbakov, 
and Bazykin, 1941; Skalon, 1951; Sludskii, 1956). 

In the Far East changes in the geographic range in the 1930's 
were evidently relatively insignificant. Along the Amur the range 
did not extend up to Gorin estuary and the lower reaches of the river. 
It encompassed only its upper reaches without reaching Lake 
Evoron and the boundary crossed the Amur roughly around Kom- 
somol. From there it turned southeast in a not-too-straight line and 
ran toward the Tatarskiy Strait at Botcha Creek and Koppa (around 
48° N; Abramov, 1954). 

Geographic range in mid-1950's. Following the extreme shrink- 
age of the boundary described above by 1930, significant increases in 
wild boar populations occurred in the 1940's and 1950's and its 
range expanded considerably. Wild boar once again began to occupy 
territory they had inhabited in the past. Part of this process com- 
menced even in the 1930's. The main reason was curtailed persecu- 
tion of the animal and in some places its protection and regulation 
of hunting. The least positive changes in the range occurred in 
Siberia east of the Altai and in the Far East. Here the present-day 
range (mid-1950's) generally corresponds to that shown above for 
the period around 1930. 

Particularly significant changes in the range in the positive sense 
occurred between the 1930's and 1950's in European USSR, Kazakh- 
stan, and parts of southwestern Siberia. Artificial shrinkage of the 
range was very intense and, obviously, the possibilities of its restora- 
tion limitless. Restoration is occurring right before our eyes as a 
matter of fact, with different stages observable in different places. In 
some sections of the range wild boar are already well restored and 
the animals present in significant numbers. Other sections are in the 
process of being inhabited but as yet do not contain a large number 
of animals. In still other sections animals are found infrequently, in 
other sections rarely, and so on. Intrusions which for relatively sta- 
ble boundaries are not significant in understanding range dynamics, 
are of great importance in the present case as an index of the ten- 
dency toward expansion and restoration of the range, and the routes 
and directions of its future limits. From this viewpoint the present 
range map of wild boar is of general interest. The boundaries given 




37 Fig. 8. Boundaries of distribution, places of transgression, and places of 

acclimatization of wild boar (Sus scrofa L.) in European USSR (scale in km). 

1 —reconstructed northern boundary of range; 2— boundary of range in 1930; 3— 
boundary of range in mid-1950's; 4— distant transgressions of wild boar after 1930— 
in the west, reckoning from the north; Rovdin region (dubious), Ulom River, Vyshnii 
Volochek, and Kamensk-Dneprov; in the east: Bashkir preserve, Agapovo at Magnito- 
gor, and Vol'sk on Volga; 5 — some places of acclimatization of wild boar between 
1950 and 1960 (Zavidovo, Serpukhov, Solotcha, and Vyksa). V.G. Heptner. 



37 

below also take into consideration the limits of intrusions (apart 
from some particularly distant ones which are specified) and provide 
an idea of the status in the mid-1950's. 

The boundary of the western habitat of wild boar in European 
USSR in the northwest proceeds initially along the reconstructed 
boundary described above, commencing at the southwestern bank of 
Lake Ladoga up to Ostashkov region and Lake Peno. From there it 
turns south toward the Nelidov region and, inexplicably, probably 
east toward the boundaries of Moscow district,^' i.e., encompassing 
the greater part of Smolensk district, probably including Vyazma, 
and proceeds into Sukhinichei region (southwest of Kaluga). From 
there, turning southeast, it proceeds towards the Lipets region, 
about 40 km south of Lipets, Voronezh (Voronezh preserve), and the 
Voronezh estuary into the Don. It then proceeds about 80 km west of 
Liski (Ukolov region) and, turning slightly southwest, to Valuek 
(Nikitov region, directly northeast of Valuek) (Melander, 1939; 
Barabash-Nikiforov, 1956; Sludskii, 1956).^^ Thus, the boundary 
encompasses from the east almost all of Smolensk district, a signifi- 
cant part of Kaluzh, possibly south of Tul'sk, the entire Orlov 
region, and western part of Lipets, Voronezh, and Kur' districts. 

The range boundary of wild boar in the Ukraine can only be 
generalized. It covers the eastern part of Kharkov district (Sludskii, 
1956) and in the south Poltava, Kiev, Kirovograd, and Vinnits dis- 
tricts (Korneev, 1952). Evidently the boundary from Valuek region 
runs southwest and encompasses the northwestern half of Kharkov 

^'Sightings in Moscow district reported recently are not taken into consideration 
here since they are associated with migrations of animals released there for acclimati- 
zation in Zavidovo (see below). In any case, even if there are some transgressions from 
the west, it is difficult to distinguish them. 

Recently, some deep intrusions of wild boar are known in the more northern part 
of the range, i.e., in the region of Vyshnoi Volochek and in the region between Lake 
Beloe and Kuban and Rybinsk reservoirs (Ulom region of Vologod district; Nasimo- 
vich, 1955). The latter falls roughly 400 km of the nearest segment of the recent 
eastern boundary. It has been suggested that these represent deep intrusions of ani- 
mals acclimatized in the southern part of Kalinin district. Also, there are cases (Kholos- 
tov, 1956) of catches of wild boar (January, 1953) even farther away in the northeast in 
Rovdin region, Arkhangel'sk district (in Vage above Shenkur, V.G.), roughly 650 to 
700 km away from nearest boundary. Evidently the reference here is to a solitary 
animal. The age of the animal, about 2.5 years, also supports this view. (V.H.) 

T"he assumption is unfounded that wild boar in Podon'e could have appeared 
not only from the west but also due to dispersal of animals released for acclimatiza- 
tion in Ryazan' district (see below) (Barabash-Nikiforov, 1956; Sludskii, 1956). The 
animals, firstly, were settled far in the north and, secondly, all of them perished. 
There is no doubt that the range expansion in Don basin is the result of animals 
spreading from western districts. 



38 

district and later turns steeply west-southwest, intersecting the 
Dnieper somewhere above Dnepropetrov and running toward the 
Dnestr evidently in Soroka district. From there, it descends along the 
river to the sea and runs along the coast into the estuary of the 
Danube River." 
40 It can be seen from the above review that over the past 20 to 25 
years the boundary moved eastward a distance of up to 250 or even 
600 km. In the northern section it has already regained its natural 
range (coinciding with the "reconstructed boundary") over a signifi- 
cant area (north of Ostashkov). This is not so for much of the range. 
The western section within the USSR continues to remain isolated 
from the rest, primarily from the Caucasian sector. 

The Caucasian part of the range from 1930 onwards did not 
expand perceptibly north, the boundary from the Azov coast to the 
Volga remaining essentially stationary. 

An extremely intense and rapid expansion of the range and a 
northward shift of the boundary occurred in the area from the Volga 
to the Altai. In the area between the Volga and Ural wild boar, 
evidently from the Caspian reeds, colonized the region of Kamysh- 
Samarsk lakes and the lower reaches of Uzen' and from there have 
spread even farther. The points at which the animals penetrated this 
region are the Bykov farmstead on the Volga (Bykov, south of Kamy- 
shin), the region of Pallasovka (east of Kamyshin), the lower reaches 
of Eruslan (Valuek region), the region between the Volga and the 
southern tributary of Eruslan-Torgun (i.e., area east of Kamyshin), 
and even Vol'sk. Somewhat more to the east wild boar have been 
sighted along the Chizhin flood zones (between Bolshoi Uzen' and 
Ural'sk) and at the Ural (Stroganov, 1954; Zaletaev, 1952; Sludskii, 
1956). The encounter at Vol'sk is an intrusion quite isolated from 
the remaining ones. It is significant that wild boar do not breed 
along Volga and the intrusion at Bykov farmstead occurred from the 
east, from the steppes, and not along the river valley from the south. 
Along Ural River the animals spread from the sea to Ural'sk. 

From the Ural the boundary turned into the lower reaches of the 
Ilek into Burannyi region and turning round the Ural and its bend 
from the south, ran into Adamovka region (Adamovka, northeast of 
Orsk). A particularly distant transgression was noticed at a place 
slightly south of Magnitogorsk (Agapovka). From Adamovka, the 
boundary ran northeast, crossed the Tobol, and running along the 
lakes lying along its right bank, somewhat east of Kustanay, turned 

"'in the Dnieper basin bodies have been sighted in river floods at Kamensk- 
Dneprov Kamensk (Sludskii, 1956). 



39 

to the north reaching Kosulin region (southwest of Kurgan at 64° E 
long, and roughly 55° N lat. — V.H.). This was the northernmost 
residence of wild boar in this part of the country. The natural range 
here had almost been restored or the animal came very close to the 
former northern boundary. 

From Kurgan the boundary turned east-southeast toward Petro- 
pavlov. From there the boundary line becomes extremely complex 
and less well defined. From Petropavlov the boundary proceeded 
directly to the south, and, having reached roughly 52°N lat., turned 
sharply east and exited at Ermentau highland (Ermentau; at 52° N 
lat. or slightly more north, directly north of Akmolinsk; see footnote 
on p. 34). From there, going round Akmolinsk in the form of an arc 
from the west, the boundary line passed through the upper reaches 
of the Nura and slightly northeast of Karaganda, encompassed Kar- 
karalin and Chingiztau mountains, and ran to the eastern tip of 
[Lake] Balkhash. From there it turned northeast, covering the Tar- 
bagatai, the Zaisan district, and the southern slopes of Kurchum 
range, and ran into China. 

Sometimes wild boar extended far beyond the limits of the above 
region into Bayan-Aul region, Maisk (at Irtysh below Semipalatin), 
into the environs of Semipalatinsk, Krasnyy Kut, at the point of 
intersection of the Irtysh and 76° E long, and even the Irtysh below 
Krasnyy Kut. Intrusions were known in Bukhtarma valley, Srosten 
pine forest in the Altai steppes (northeast of Rubtsov), toward the 
confluence of Biya and Katun, and in other places of pre-Altai 
steppes. 

As can be seen from the data presented thus far, in the past 20 to 
25 years (commencing from 1930) the habitat of wild boar between 
the Volga and Altai has expanded sharply. In Kazakhstan expansion 
was particularly intense from 1941 . From the beginning of the war a 
sharp reduction in hunting of wild boar took place, and a humid 
period set in again so that all the reservoirs which had dried up 
41 became favorable habitats once more for animal life; they filled with 
water and various types of reeds and cattails grew on the shores. 
These factors and subsequent regulation of hunting led to an expan- 
sion of the range of wild boar. 

Over the last 20 years the habitat of wild boar has tripled in 
Kazakhstan, covering territory where it was absent 100 to 150 years 
ago. It has colonized once again not only the steppe but the forest- 
steppe zone. In the western and central part of the area under de- 
scription the present-day northern boundary of the range has already 
come close to the former natural ("reconstructed") boundary and at 
one place even reached it. In the east, as shown By intrusions (see 




he 



о л О 



"о 

> 



н 5 



-3 I -^ 



л ^ 



^ — . :; 



41 

map), the very same process has occurred, although it has not 
attained the same intensity. 

The rate of colonization of wild boar, especially in the 1940's, 
was extremely high at some places. Over a period of two to five years 
in some parts of Kazakhstan wild boar have spread 300 to 500 km (in 
a straight line), moving roughly at an average of 100 km each year. 
Colonization has proceeded mainly along river valleys and lake 
systems. 

On the whole the expansion and reconstruction of the range of 
wild boar in the area under description was more significant than in 
the west. This is simply explained by the low population density in 
Kazakhstan and adjoining places. ^^ 

The expansion of this part of the range led to its merger in the 
Soviet Union with the Caucasian sector. However, the separation of 
eastern and western parts of the range is still total and very distinct. 
There is no possibility, at least not in the near future, of the two 
ranges merging, but the eventuality cannot be totally ruled out. A 
tendency to merge from both sides is discernible. 

As for the areas of distribution of wild boar in southern Altai 
(only stray occurrences), in Tuva, the Sayans and farther east, appar- 
ently no tendency for expansion and restoration of the range has 
been detected in the last decade. 

For purposes of acclimatization wild boar were released for hunt- 
ing in the last decade in Serpukhov andZavidov near Volzh reservoir 
northwest of Klin (Kalininsk district), Solotcha region on the left 
bank of the Oka slightly north of Ryazan' (Ryazan' district), and in 
Vyksa region on the right bank of the Oka slightly south of Murom 
(Arzamas district). Zavidov wild boar survived until 1955 and settled 
to some extent especially in Klin region of Moscow district. It is 
possible that it was from here that stray intrusions were reported in 
Vyahnei Volochek and Ulom region. Solotchin wild boar, left to 
their fate in a highly snowy district (Zavidov boars survived), died 
out completely by 1950's.^' 



It is premature to assume that wild boar in Kazakhstan has "completely" 
regained its former geographic range (Sludskii, 1956). 

'Propagation of wild boar between the Volga and Altai cited from data of Sluds- 
kii (1956). 

As a result of cultivation of virgin land the situation may undergo intense and 
rapid changes. 

The suggestion that animals from here were seen in the upper reaches of the Don 
(Sludskii, 1956) has no basis (see above). 



42 

Vyksa wild boars were still surviving in the early 1950's but 
evidently are condemned to die. 

Geographic Range outside the Soviet Union 

The extensive range outside the Soviet Union has been shown in 
Fig. 4. In Africa it includes Rio de Oro (Spanish Sahara), Morocco, 
Algeria, and probably Tunisia (at present, however, wild boar are 
absent there), Egypt (disappeared around 1900), and Sudan (Kordo- 
fan and Sennar). In Europe the range covers all western Europe 
including the British Isles (extinct), Corsica, Sardinia, and Sicily. In 
the north the range is bounded by the Baltic Sea but also covers the 
southern part of the Scandinavian Peninsula (extinct). The Asian 
range of wild boar includes Asia Minor, Syrian zone of United Arab 
43 Republic and Palestine, Iran, Afghanistan, India, Baluchistan, 
Kashmir, Nepal, Ceylon, Burma, Indochina, Malacca, Andaman 
and Nicobar Islands, Sumatra and adjacent tiny islets, Java and the 
entire chain of Sunda islets in the east including Flores, Sumba, 
Banks, Sumbawa, Timor, and Rota. Farther away the range covers 
Aru and Kai, New Guinea, Louisiade, Solomon Islands, Bismarck 
Islands, Admiralty Archipelago, and some islands on the western 
part of New Guinea — Yapen, Misol, Barenta, and others, including 
Vaigeo and Seram Islands. In the north the range covers Hainan, 
Taiwan, and all the islands of Japan, including Hokkaido and 
Ryukyu. 

On the mainland the range covers northeastern, eastern, and 
southeastern China far south. In the west, in this part of China, the 
range covers Szechwan, Shensi, and Kansu almost up to Kukunor. 
In Mongolia wild boar occur in Hentei roughly up to 47° N lat. to 
the south, Sayan and Altai region, Khangay in the south roughly up 
to 46° N lat., and the Mongolian Altai to the south up to 44° N lat. 
In northwest China the range includes the Tien Shan system, Tarim 
and Yarkand basins, and extends toward the mountain system of 
Dzhungaria district. In Inner Mongolia, in the desert region of 
northwestern China and Tibet, wild boar are absent and were never 
present. Wild boar are already extinct in several places apart from 
those indicated above. 

Wild boar are either acclimatized for hunting or have become 
feral (Chinese domestic swine) in some places in the United States 
(Tennessee, North Carolina, Georgia, Texas, California, New 
Hampshire) and in several places in Central America. They have 
similarly acclimatized at two places in Argentina and settled there 
rather rapidly. (V.H.) 



43 

Geographic Variation 

The geographic variation of the species in the area under discus- 
sion is very high and 26 to 28 forms are recognized. 

The number of varieties is extremely large, exceeding 50. The 
actual number of subspecies, however, is undoubtedly less than 26 to 
28, since a large number of descriptions are unsatisfactory and the 
intraspecific variation of the species has never been revised. 

In the Soviet Union there are five subspecies which differ slightly 
in some details of craniology, mean body and skull size, and domi- 
nant color type.^^ 

1. Central European wild boar (S. s. scrofa Linn., 1758) (syn. falz- 
jeini). 

Size, medium. Maximum skull length 370 to 410 mm. Color dark 
and fur rusty-brown. Lacrimal long and relatively narrow. Third 
upper molar relatively long; talon well developed and partly forms 
third transverse row of tuberosity. 

Found in west European USSR. Outside the USSR found in 
central and western Europe. Relationship with other subspecies in 
the context of its range and some other characteristics not clear. 

2. Rumanian wild boar (S. s. attila Thos., 1912). 

Size, large. Maximum skull length 410 to 470 mm. Color dark 
but lighter than that of preceding form. Lacrimal long. Third upper 
molar same as in preceding form. 

Found in the Caucasus, Trans-Caucasus, and along northern 
Caspian coast. In some respects wild boar of the Ukraine and eastern 
part of Belorussia and probably of southwest Turkmenia also 
belong to this form. Outside the USSR, apart from Rumania, also 
evidently found in Hungary, the Balkans, Asia Minor, and northern 
Iran. Relationship to S. s. lybicus Gray, described from southwest 
Asia Minor, also not clear. 

3. Middle Asian wild boar (S. s. nigripes Blanf., 1875). 

Size, generally quite large but range of variation quite high. 
44 Color light, lighter than rest of subspecies, and stands out in con- 
trast to black color of legs. Fur lighter than in western forms. 
Lacrimal shorter than that of preceding forms, and facial portions 
of skull somewhat shortened. Third upper molar as in European 
forms but sometimes simpler. 

^^According to Adlerberg (1930) and Fetisov (1956), with modifications. 



44 

Found in Middle Asia and Kazakhstan. Outside the USSR found 
in eastern Tien Shan, western Mongolia, Kashgar, and possibly 
Afghanistan and southern Iran. 

4. Trans-Baikal wild boar (S. s. sibiricus Staffe, 1922) (syn. raddeanus). 

Smallest of wild boar forms. Maximum skull length of male 370 
to 400 mm. Color dark brown, almost black. From posterior edge of 
mouth, along cheek to pinna, gray or grayish-whitish patch up to 24 
cm in length occurs. Skull shortened and high; lacrimal almost 
squarish. Third upper molar short and relatively broad; talon 
poorly developed and simpler than in European forms. 

Found in Baikal and Trans-Baikal. Outside the USSR found in 
northern and northeastern Mongolia. Relationship of this form to 
the form canescens Heude, described from near Peking, is not clear; 
Manchurian wild boar are not related to this form. 

5. Ussuri wild boar (S. s. ussuricus Heude, 1888) (syn. continentalis, 
gigas). 

Largest of USSR swine and of species as a whole. Maximum 
length of male skull 490 to 510 mm. Color variable but mainly dark; 
new coat dark brown or almost black. From corner of mouth to ear 
distinct white band present (G.F. Bromlei). Lacrimal somewhat 
shortened but longer than in Trans-Baikal form. Third upper molar 
broad. 

Found in Ussuri basin and Amur in the west to Great Khingan. 
Evidently this form inhabits northeastern China and the forms de- 
scribed here, songaricus Heude (Sungari valley) and mandchuricus 
Heude (Mukden), must also be placed as its synonyms. 

The number of wild boar forms described from parts of the range 
falling outside the USSR is very large, more than 20. This number is 
made up of not only the subspecies, which are geographically and 
morphologically close to the group scrofa, but also those which 
were formerly placed among other species but are now merged with 
the species S. scrofa (see above). With such an interpretation of the 
species, the morphological range of its geographic variability is con- 
siderable. Along with the giant forms of Europe, Amur region, and 
Manchuria, in the far south and especially in the island region of the 
extreme southeastern part of the range, there also exist small, often 
extremely small, forms. The extreme form is represented by Papuan 
swine which are no larger than a young pig. 

Not all the forms have been described with adequate supporting 
information and there is no doubt that their actual number is fewer 
than currently asserted. Some forms of the island region of southeast 



45 

Asia evidently represent domestic swine turned wild, which were 
imported by islanders at various times from different islands. 

At present the following subspecies of S. scrofa (Haltenorth and 
Trenze, 1956) are commonly recognized in Europe (in addition to 
forms scrofa, attila, and falz-feini — see above): S. s. meridionalis 
For s.- Major, 1882 (Sardinia), S. s. castillanus Thos., 1912 (Iberian 
Peninsula), S. s. reiser i Bolkay, 1925 (Yugoslavia), and S. s. major i 
de-Beaux and Festa, 1927 (Italy); in Africa: S. s. barbarus ScL, 1860 
(northwest Africa, Rio de Oro, Egyptian region of United Arab 
Republic and Kordofan); in Asia (in addition to attila, ussuricus, 
sibiricus, and nigripes- — see above): S. 5. libycus Gray, 1868 (south- 
ern Turkey and Palestine), S. s. cristatus Wagn., 1839 (India, Indo- 
china, Burma, and Ceylon), S. 5. vittatus Miill. and Schleg., 1842 
(Sumatra, Java, Bali, Lombok, Sumba, Banka, and Sumbawa), S. s. 
floresianus Jent., 1915 (Flores), S. s. timorensis Mull, and Schleg., 
1845 (Timor), S. s. nicobaricus Mill., 1902 (Nicobar Island), S. s. 
andamanensis Blyth., 1858 (Andaman Islands), S. s. chirodontus 
Heude, 1888 (southern China and Hainan), S. s. taivanus Swinhoe, 
1868 (Taiwan), S. s. riukianus Kuroda, 1924 (southern islands of 
45 Ryukyu archipelago), S. s. leucomystax Temm., 1842 (Japan), S. s. 
koreanus Heude, 1897 (Korean peninsula), S. s. moupinensis M.-E., 
1871 (from Szechwan through Central China to Jehol), and S. s. 
papuensis Less, and Gerot, 1826 (extreme southeastern part of the 
range apart from the islands indicated above, with the center in New 
Guinea). (V.H.) 

Biology 

Population. The wild boar jDopulation has undergone sharp 
changes in different years. Fluctuations are determined in each indi- 
vidual district by litter size, migrations, mortality in severe winters, 
occurrence of infectious diseases, and intensity of hunting. In just a 
year or two a population can drop over a considerable area by 10%, 
doubling or trebling in the very next year (even without migrants). 
Precise data on wild population and density in the USSR are 
few. The total number of animals in the Soviet Union is evidently a 
few hundred thousand. In Belovezh Forest in the 1890's and early 
twentieth century the population density ranged from 4.4 to 30.8, an 
average of 13.3 per 1,000 hectares (Kartsev, 1903); in the 1940's and 
1950's it fluctuated between 4 and 19 (average 9.9) (Lebedeva, 1956). 
In the Astrakhan' preserve at the end of the 1930's the population 
density rose to 27 per 1,000 hectares (Lavrovskii, 1955), in the Cauca- 
sian preserve averaged 4 to 8 (Donaurov and Teplov, 1938), and in 



46 




Fig. 10. Broad-leaved forest, typical habitat of wild boar in Belovezh Forest. 
Photograph by A.G. Bannikov. 



Bryansk district forests 0.5 to 4.2 (Fedosov and Nikitin, 1951). In 
years of abundant pine nut yield in Khentei (Mongolia), and evi- 
dently also in the Far East, population density rose to 30 to 40 per 
1,000 hectares. In Kazakhstan and Central Asia along the banks of 
large lakes and rivers densities of up to 50 to 60 per 1 ,000 hectares are 
known in favorable years. 
46 As a result of sporadic migrations, for example, during forest 
fires or heavy snows, temporary concentrations of animals up to 200 
to 300 per 1,000 hectares form in reed thickets along the banks of 
lakes over a few hundred hectares. 

Throughout the eighteenth and the first half of the nineteenth 
century a reduction in population and range of wild boar took 
place. Population growth was recorded from the second half of the 
nineteenth century in Baltic region (Ling, 1955) and from the 1930's 
everywhere else (with the exception of some regions of the Caucasus, 
Trans-Carpathia, and Siberia). Restoration of the range occurred 

^'in Germany at the end of the last century the population density of wild boar 
was 1.7 per 1,000 hectares; between 1936 and 1939 it rose to 5.2 when the total popula- 
tion was 66,000 (Oloff, 1951). 



47 

simultaneously (see above). Maximum population of wild boar was 
observed in the 1950's in the Volga delta, Syr-Darya and Chuya 
valleys, lower reaches of the Hi River, along the banks of Lake Zai- 
san, and in western Tien Shan mountains (Sludskii, 1956). In the 
Far East it was high along midreaches of Khor and Anyuya Rivers, 
along the Shukhi-Pokta range, along the northern spurs of the Lit- 
tle Khingan, on Iman and Bikin Rivers, and at the sources of the 
Bidzhan River (Abramov, 1954). 

Before the commei/cement of the nineteenth century many wild 
boar lived in downstream valleys and reed thickets of steppe rivers 
(Dnestr, Dnieper, Don, and Danube). In the deltas of the Kuban, 
Volga, lower reaches of Kuma, Terek, and Kur' Rivers and along the 
coasts of the Caspian Sea wild boar are numerous even today. 

Habitat. The diversity of diet of wild boar determines its extreme 
eury topic character. It inhabits zones from boreal taigas to deserts. 
In mountains it may be found in all zones up to and including the 
alpine. In the Carpathians wild boar occur up to 1,900 m, in the 
Caucasus up to 2,600 m, and in the mountains of Central Asia and 
Kazakhstan up to 3,600 to 4,000 m. The importance of a given loca- 
tion for the species is determined by the abundance of food, its 
accessibility, and the protective conditions offered. With changes in 
seasonal and annual abundance of food sources, the importance of a 
place also changes. Nevertheless, some major locales are usually 
occupied by wild boar for most of the year, since they ensure food 
and offer hideouts to the animal. Other places are important only 
temporarily and still other areas always of secondary importance, 
being visited only periodically and by a small number of animals. In 
Europe deciduous and mixed forests currently serve as the main 
habitats of wild boar. Forests in which most of the trees are oak or 
beech and which enclose marshy sections and meadows serve as 
optimum habitats. Sites used periodically include meadows and cul- 
tivated lands. Thus in Belovezh Forest, in woods and in open sites 
85.1% and 14.9% respectively of wild boar were counted over a period 
of four years. The main habitats in the region are represented by 
well-developed, broad-leaved, mixed forests and marshy mixed 
47 forests. Periodically, in spring and autumn, meadows and farms are 
also used. Coniferous forests and undergrowths are of secondary 
importance (Lebedeva, 1956). Pure oak groves and beech forests 
acquire importance as habitats of wild boar only in those years when 
these species bear fruit. 

In the montane regions of the Caucasus and Trans-Caucasus 
belts of beech and oak and other fruit-bearing forests are of primary 
importance for wild boar, which aie found in them year-round (Din- 



48 




46 Fig. 11. Marshy forest, a favorite habitat of wild boar in Belovezh Grove. 

Photograph by A.G. Bannikov. 



nik, 1910; Loginov, 1936; Donaurov and Teplov, 1938; Nasimovich, 
1939; Markov, 1932, 1933, and 1934). In these forests, 37 to 62% of all 
the wild boar counted were found in the Caucasian preserve. Coni- 
ferous forests (8 to 55%) and subalpine zones (2 to 28%) play a second- 
ary role, where wild boar usually live from July to September. The 
alpine zone is used by wild boar only occasionally during the five 
summer months and the occurrence rate is low (Donaurov and 
Teplov, 1938). 

In the mountains of Middle Asia and Kazakhstan wild boar col- 
onize all vertical zones. The main habitats are the lower parts of fir 
and deciduous forest belts. The latter acquire particularly great impor- 
tance in the period of ripening of fruits and nuts. Tugais [vegetation- 
covered bottomlands] along mountain rivers and juniper forests 



49 



48 




Fig. 12. Reed thickets on the Volga delta, typical habitat of wild boar. 
Photograph by A.G. Bannikov. 

are important locales for wild boar in different periods of the year. 
Subalpine meadows are visited by boar in summer and the steppe 
slopes in spring and autumn (Antipin, 1941; Flerov, 1935; Zverev 
and Keleinikov, 1947; Shul'pin, 1948; Ivanenko, 1956). On the low 
mountains of the Kazakh hillock zone (Chingistau, Karkaralin hills, 
and others) wild boar live mainly in marshy valleys overgrown with 
aspen, birch, rose, etc. at a height of 500 to 800 m (Sludskii, 1956). 

In the mountains of the Far East (Sikhote-AIin) the main habi- 
tats of wild boar are nutpine groves, hilly mixed forests with Mon- 
golian oak and Korean nutpine, swampy mixed taiga, and coastal 
oak forests. Now and then, in small numbers, boar invade the subal- 
pine zone with dwarf nutpine. The distribution of the animal varies 
in different seasons and is particularly dramatic in years of failure of 
major food crops (Abramov, 1954; Rakov, 1956). In the mountains of 
Trans-Baikal wild boar are confined to nutpine and shrubs in the 
river valleys (Kopylov, 1950). 

In the plains of the Trans-Caucasus, Kazakhstan, and Middle 
Asia the main and most important habitats of wild boar are thickets 
of reed, cattails, and shrubs along river and lake banks. Where large 
areas of such shrubs are available wild boar generally occupy them 
year-round. 



50 



Dense shrubs along river banks consisting of willows, rose, 
honeysuckle, oleaster, etc., and especially swampy forests can be 
considered permanent but secondary habitats of these animals since 
they primarily seek cover in them. Wild boar also live permanently 
in marshy birch-aspen forests among strips of pine groves in eastern 
Kazakhstan. Sand knolls and short, prominent hillocks sporting 
undershrubs also periodically serve as secondary habitats. In 
Kazakhstan and Central Asia wild boar also visit saxaul woods and 
groves oiNitraria (Dinnik, 1910; Flerov, 1935; Shul'pin, 1948; Lav- 
rovskii, 1955; Sludskii, 1956). In some regions of Tadzhikistan and 
Badkhyz (southern Turkmenia) boar are regularly encountered in 
pistachio groves in winter; in spring they fan out great distances into 
open desert, feeding on meadow grasses, sedges, and tulips 
(Heptner, 1956). In every case wild boar periodically feed on culti- 
vated crops, especially when crop failures occur in their natural diet. 




Fig. 13. Birch forest zone in the Caucasian preserve, habitat of 
wild boar. Photograph by A.G. Bannikov. 



51 




49 Fig. 14. Subalpine meadow in the Caucasian preserve. Furrows made by wild 
boars. Photograph by L.S. Lebedeva. 

Food. A comprehensive list of the diet of wild boar would be 
extremely long and include products of vegetable as well as animal 
origin. The main items can be divided into four groups: 1) rhizomes, 
roots, tubers, and bulbs of plants dug up by the animal throughout 
most of the year and throughout the entire range; 2) products of 
fruit-bearing trees, nuts, berries, and seeds which the animal eats 
after ripening, digging them up from under the snow in winter 
when the crop is abundant; 3) aerial parts of plants; green parts of 
herbaceous plants are used in the period of their availability, while 
bark, twigs, and shoots often constitute compulsory food, along 
with rubbish, in winter; and 4) items of animal origin — earthworms, 
insects and their larvae, mollusks, fish, rodents, insectivores, bird 
eggs, lizards, snakes, frogs, carrion, etc.; these serve as food more 
often in the warm period of the year but also year-round to some 
extent. 
49 Food which wild boar extract from the soil (underground plant 
parts and animal food) constitutes a greater portion, (in terms of 
quantity) of their diet than food gathered above the soil. In Belovezh 
Forest boar dig up underground food which in weight is 3.5 times 
(average) more than that gathered above the soil (Lebedeva, 1956). 
Digging up the soil to gather food is highly characteristic of wild 
boar. 



The plains of Kazakhstan and eastern Siberia where underground food is less 
accessible to wild boar in winter because the top soil freezes constitute an exception. 



52 

In different parts of an extensive range the diet composition of 
wild boar differs markedly both in terms of category of food items 
and its species composition. 

In western Europe underground plant foods include roots of 
bracken {Pteridium aquilinum), willow herb, bulbs and roots of 
meadow herbs, and bulbs of cultivated crops. Wild boar gather this 
food in early spring and summer and when acorn and beechnut 
crops fall short, in winter and autumn also (Kissling, 1925; Sneth- 
lage, 1934; Oloff, 1951). Acorns, beechnuts, chestnuts, hazel nuts, 
and more rarely fruits of fruit-bearing trees serve as excellent food. 
Wild boar invariably prefer beechnuts but acorns, when abundantly 
available, serve as the most important "fattening" food (Oloff, 
1951). A small quantity of the green parts of meadow and marsh 
grasses and other graminaceous plants is consumed by wild boar in 
April and May. 

Food of animal origin is extremely diverse. It includes earth- 
worms and various insects and their larvae gathered by the animals 
from the forest bed, including such forest pests as June bugs and 
pine geometrid moths (Fitingof-Rish, 1952; Klemm, 1944; Khaber, 
1950; Tsappe, 1958; Yanda, 1958; and others). In years of abundant 
availability rodents also constitute a significant food item for wild 
boar (Oloff, 1951). Other animal foods such as lizards, mollusks, 
snakes, bird eggs, etc., represent occasional components in the diet 
of this animal. 

The food of wild boar has been studied in detail for the European 
part of the USSR only in Belovezh Forest (Severtsov and Sablina, 
1953; Sablina, 1955; Lebedeva, 1956). The list of wild boar food in 
50 Belovezh Forest contains 118 items of plant and animal origin. The 
underground food (39 plant and 11 animal items ) and aerial parts 
of plants (42 species) are most diverse. Underground plant parts 
(roots and rhizomes) constitute the most regular food items and are 
consumed year-round in large quantities. Maximum quantities of 
root foods are gathered in April (on the average 404 g per 1 m') and 
August (418 g) and minimum amounts in July (237.2 g). In this 
group of food wood anemone, knotweed, and saltwort [Polygona- 
tum] are of utmost importance; in some periods of the year butter- 
cup, calla, dandelion, and bennet play an important role. 

Acorns and hazel nuts are occasional foods. This is because years 
with good yields are few among years when oak and hazel do not 
bear fruit. Hazel nuts with their poor and rare yield are insignificant 

^'The number of species is more than 39 and 11; root and animal foods are often 
identified only up to genus. 



53 




Winter Early spring Spring- Summer Summer Early Autumn 

PERIODS 

Fig. 15. Variations in food of wild boar in Belovezh Forest around the year 
(from Lebedeva, 1956). 

as food of wild boar. In years of good acorn yields the animal con- 
sumes them avidly throughout all of autumn and winter, gathering 
up to 150 g of acorns from 1 m . But since crops of acorn appear only 
once in three or four years this food is generally not of primary 
importance to wild boar in Belovezh Forest. 

The green parts of plants represent a seasonal food consumed in 
May to June when wild boar gather an average of up to 1 19.2 g per 1 
m^. Of the plants from which wild boar eat the green parts five 
species are primary. Marshy thistle with its large succulent leaves 
occupies first place, sedge second place (consumed in May when the 
leaves have not yet become coarse), and nettle, touch-me-not, and 
bennet come third. The vegetative portions of plants are also con- 
sumed by wild boar in winter and horsetail, bilberry, heath, and 
herbaceous twigs figure prominently. The animal uses these foods 
intensely in years when the acorn yield is inadequate due to soil 
freezing. 

Among animal foods earthworms are of utmost importance in 
the diet of Belovezh Forest wild boar. In the total number of cases in 
which animal foods were recorded earthworms constituted 62%, 
insect larvae 31.9%, and the remaining 6.1% other types of animals. 
Quantitatively earthworms represented 74%. Animal components 
fall in the category of regular food but play a vital role in the warm 
season of the year when wild boar gather an average of 101 g earth- 
worms from 1 m^. In the winter, because of their poor accessibility, 
earthworms are less significant. 

Thus in Belovezh Forest wild boar gather much of their food 
from the soil and forest beds. By overturning the soil with their 



Average 
for the 
year 



54 

muzzle they collect fruits and seeds of trees and shrubs and also the 
larvae of some insects. Rooting through the soil layers they gather 
51 earthworms, larvae of insects, and underground plant parts. Bur- 
rowing fairly deep pits in search of food sources they uncover 
rodents or rootstocks of marsh plants. Boars dig year-round but 
mostly in winter and early spring when the maximum amount of 
underground food is utilized. On the average in one year a single 
boar gathers 1,490.8 g underground food and 452.2 g aerial plant 
food from 1 m^ (Lebedeva, 1956). 

Apart from the foods mentioned above Belovezh Forest wild boar 
also consume potatoes, artichoke roots, and cereal crops. However, 
only part of the population penetrates farms and that, too, mainly in 
years of failure of forest food crops. Mushrooms and rodents are of 
minor importance in the food of wild boar (Severtsov and Sablina, 
1953; Lebedeva, 1956). 

In the montane forests of the Caucasus the main groups of food 
of wild boar are the same as those in western Europe but their role 
differs somewhat. The importance of different foods in the Cauca- 
sian preserve, based on the frequency of their occurrence in 35 dis- 
sected stomachs, is as follows: roots and rhizomes of herbaceous 
plants (65.7%), beechnuts (62.9%), green parts of herbaceous plants 
(48.6%), acorns (22.9%), vertebrates (14.3%), earthworms (8.4%), and 
pears (5.7%); insects, apples, and damson plums were found in an 
equal number of stomachs (2.9%) (Donaurov and Teplov, 1938). 

The underground parts of herbaceous plants represent perman- 
ent year-round food of wild boar but do not occupy an important 
place in their diet. These foods acquire utmost importance at the 
end of spring and early summer. They become vital in winter in 
those years when the yield of nuts is unsatisfactory. Otherwise, even 
rootstock occupies a prominent place in winter in the food of wild 
boar since the accessibility of fruits and nuts is minimal and their 
reserves exhausted (Nasimovich, 1939). In the Caucasian preserve 
the rhizomes, roots, and bulbs of eight species of plants are con- 
sumed avidly (Loginov, 1936). Donaurov and Teplov (1938) have 
listed 28 species of plants whose underground parts are consumed by 
wild boar. In high mountain meadows alone 23 species have been 
identified, the rhizomes, roots, and bulbs of which are consumed by 
these animals (Lebedeva, 1956). Evidently the list is much longer 
than generally supposed. 

The fruits of nut-bearing trees, wherever available in the Cauca- 
sus and Trans-Caucasus, represent the most important food item for 
wild boar during most of the year when their yield is abundant. The 
fruits of fruit-bearing trees are a seasonal food consumed for two or 



55 

three months in a year. In June and July cherries ripen and wild 
boar consume them avidly but their overall importance is only 
secondary. In August and early September the animals concentrate 
at places where damson plums fall and feed on them extensively, but 
only for a short period. Even in September, when the damson plum 
ripens, wild boar leave these thickets to feed on pears and apples 
ripening immediately thereafter. Furthermore, acorns drop in mid- 
September, a favorite food item. In November boar shift to ripening 
beechnuts and also gather fallen walnuts and chestnuts. Thus the 
fruits of nut-bearing trees and beechnuts constitute the main food of 
wild boar from October through March, i.e., for five or six months. 
In the years of their abundance these foods are well preserved under 
snow and at places where they have not been consumed in winter 
serve as an important food in spring, i.e., in March and April. 
Should wild boar come across them later in the year they will con- 
sume them even when they have sprouted. 

The green parts of herbaceous plants are of great importance at 
the end of spring and summer but are utilized in winter as well when 
wild boar consume a fairly large amount of winter-green plant spe- 
cies. In winters with heavy snows and poor yields from nut-bearing 
trees the animals consume the bark of trees and fungi (Donaurov 
and Teplov, 1938). 

Among animal foods of utmost importance to wild boar in the 
Caucasus are earthworms. The animal will root through large areas 
of meadows in search of them. It consumes a small quantity of 
52 insects, rodents, and bird eggs. Carrion is also consumed; under 
conditions of starvation instances of cannibalism have even been 
reported. Cultivated plants represent seasonal foods which play an 
important role only in some regions of the Caucasus and Trans- 
Caucasus from July to October (Dinnik, 1910; Satunin, 1906; 
Markov, 1932; Donaurov and Teplov, 1938; and others). 

In the forests of the Far East underground parts of plants also 
constitute a considerable portion of the diet of wild boar throughout 
the year. They are evidently of great importance in autumn and in 
the first half of winter, especially in the event of crop failure among 
nut-bearing trees. The animals dig up the roots and rhizomes of 
such plants as lespedeza, mountain ash, spirea, rhododendron, etc. 
In summer wild boar avidly root cow parsnip, angelica, burnet, lily, 
pea, coltsfoot, and other plants. 

Seeds of Korean nutpine and acorns of Mongolian oak are the 
most important food items. In the season of these crops, commenc- 
ing from September, wild boar feed on them in nutpine and oak 
forests until midwinter. When the crop is abundant, they will con- 



56 

tinue to consume pine seeds and acorns throughout winter and 
spring right up to June of the following year. At the end of summer 
berries are consumed (Abramov, 1954; Rakov, 1956). 

Among plants from which the aerial parts are consumed by wild 
boar in the Far East, winter horsetail is of primary importance, espe- 
cially in river valleys when nut crops are inadequate. In places 
where horsetail is not available the animals consume twigs, moun- 
tain cranberries, and terminal branches of white-barked fir and its 
needles, especially in the latter half of winter. In spring and early 
summer the green portions of such plants as nettles and some umbel- 
liferous and Amur vines constitute not less than one-half of all the 
food of wild boar. 

Throughout the warm season of the year foods of animal origin 
become essential. In summer the animals gather earthworms, mol- 
lusks, insects and their larvae, rodents, frogs and their spawn, eggs 
and nestling of birds on the ground, etc. In coastal regions wild boar 
sometimes feed on marine animals such as crabs, fish, and carcasses 
of marine animals. After salmon have completed spawning boar 
will feed on their carcasses (Siberian salmon, humpback salmon, 
etc.). 

In years of poor nut yields boar visit potato, oat, and maize farms 
at the end of summer and autumn (V.D. Shamykin; Abramov, 1954; 
Rakov, 1956). 

In the montane taiga of Trans-Baikal and northern Mongolia 
rhizomes, roots, and bulbs (especially of peonies and lilies) represent 
the most important food items for wild boar year-round on exposed 
slopes, glades, and forest edges. The animal avidly seeks out the food 
caches of voles which contain large amounts of root-stock. 

Pine seeds constitute a highly nutritional and valuable food. 
When the crop is abundant boar wholly survive on these nutritive 
cones, gathering them in autumn and digging them up from under 
snow in winter. Unfortunately nutpine yields seeds only once every 
three or four years and hence the importance of underground plant 
parts as reserve food is paramount. Pine cones are consumed whole 
by wild boar and stomach contents have yielded up to 6 to 7 kg of 
this item alone (Bannikov, 1954). 

In the spring with the onset of verdure, boar mainly consume 
whatever green vegetation abounds. In winter, when verdure is 
scarce, mushrooms, fungi, twigs, cranberries, and pine needles 
assume considerable importance. 

Among foods of animal origin which boar gather in the warm 
period of the year, earthworms and rodents play an important role. 
The latter, i.e., forest and narrow-skulled voles, are found in large 



57 

numbers in the stomachs of boar in winter (Bannikov, 1954; 
Kopylov, 1950). 

In the mountains of Kazakhstan and Middle Asia underground 
plant parts represent a constant and essential food for wild boar. In 
the Trans-Ili Alatau the animals feed on bulbs, rhizomes or roots of 
geranium, tulip, pea, lady's mantle, dandelion, and onions year- 
round (24 to 100% of finds). These foods begin to predominate in 
September and become the main food in winter up through March. 
53 On Dzhambul mountain (Bet-pak-dala) even at the end of May boar 
mainly survive on tulip bulbs and rootstock of Oliver's gentians 
(Sludskii, 1956). In the Vakhsh range (Tadzhikistan) underground 
parts of 18 plant species have been found in the stomachs of 93 
trapped wild boar. From December through March bulbs and root- 
stocks greatly predominate in the stomach contents of wild boar; in 
summer and autumn underground parts of plants are always found 
in the stomachs but in small quantities (Ivanenko, 1956). 

Fruits are important seasonal food of wild boar in most montane 
regions of Kazakhstan and Middle Asia. On the northern slopes of 
the Trans-Ili Alatau and Karatau wild boar feed on fallen apples in 
autumn. Boar of the Talasski Alatau and Pskem and Ugam ranges, 
as well as in other mountains of Middle Asia, gather fruits of wal- 
nuts, damson plums, plums, and pistachios (Sludskii, 1956). On 
Vakhsh range fruits of damson plums, hawthorn, dog rose, and 
walnut, pods of flax, and ears of wheat are predominant foods of 
wild boar in August through September (Ivanenko, 1956). In west- 
ern Kopet-Dag the usual food comprises pistachios, damson plums, 
figs, blackberries, almonds, walnuts, pears, and other fruits. In 
Uzbekistan and Kirgizia almonds, walnuts, pistachios, and grapes 
are excellent fattening foods (Kashkarov, 1932; Flerov, 1935; and 
others). 

In the Trans-Ili Alatau from the end of April through August the 
aerial parts of plants, mainly herbage, predominate (40 to 53% of 
finds). They are also found in small quantities from January to 
March as well as in September and October (Sludskii, 1956). Parts of 
15 plant species have been found in the stomachs of wild boar 
caught on the Vakhsh range; in April and June boar consume leaves 
and stalks of grasses along with seeds of leguminous plants (Iva- 
nenko, 1956). Foods of animal origin found in the Trans-Ili Alatau 
boar comprise larvae of insects, worms, terrestrial mollusks, and 
rodents. The latter, when prolific in October, constitute an appreci- 
able percentage of boar food (Sludskii, 1956). 

On the Volga delta and along other rivers of the Caspian, and 
rivers and lakes in the plains of Kazakhstan and Middle Asia, the 



58 

main foods of boar year-round are rootstocks of reeds and cattails. 
This type of food invariably predominates and its importance drops 
somewhat only in the period of high floods in May and June, in the 
event of severe freezing of soil in winter, or for a short period when 
highly nutritive foods such as fruits of water caltrops or locusts are 
available. In addition to the above-named primary plants, the 
underground parts of which are consumed by wild boar, rootstocks 
of canes, roots of reed grass and sedge, and bulbs of sagittaria are 
also dug up by the animal. In some places boar avidly dig up roots 
of licorice, Indian hemp, spurges, thistles, and other plants. 

In the deserts between the Caspian Sea and Lake Zaisan in 
summer, especially from the middle of April to July, ferula roots 
assume much importance. In search of these roots wild boar leave 
river catchment areas and wander many kilometers deep into the 
desert. Sometimes the animals feed on starchy ferula roots in 
autumn and early winter as well. In deserts boar dig up roots of tiny 
saltwort and saxaul [Haloxylon] (Karelin, 1875; Lavrovskii, 1955; 
Sludskii, 1956; and others). In the Badkhyz (southern Turkmenia) 
wild boar enter the open desert from the mountainous parts of the 
country (Gyaz-Gyadyk) in winter and spring and feed not only on 
the aerial parts of plants (mainly meadow grass and sandy sedge 
forming dense thickets in some years), but also on their under- 
ground parts and on bulbs of numerous tulips. The latter, evidently, 
is a favorite food of wild boar (Heptner, 1956). 

Fruits and seeds are seasonal foods of wild boar of the desert- 
steppe regions and important only in some regions. In years when 
water chestnuts are available wild boar of the Volga delta and Lake 
Zaisan consume them avidly. Having begun to consume these 
highly nutritive fruits from August, the animals become extremely 
54 fat by the time of freezing weather (Yablonskii, 1875; Lavrovskii, 
1955). In the Hi River delta and in several other places in July and 
August berries (drupes) oi N itraria assume great importance and are 
consumed by wild boar directly from the bush; the animals wade 
deep into salt marshes for this purpose. At the same time, in tugais, 
they avidly consume blackberries and, from the beginning of Sep- 
tember, oleaster fruits (Sludskii, 1956). In some places they forage in 
fields of millet, barley, wheat, and also melon fields. 

Among plants, the vegetative parts of which play a significant 
role in the diet of wild boar of this region, reed shoots are important. 

"in China (formerly, probably also in the south of the Soviet Far East) fruits of 
water chestnuts and the rosettes of lotus with seeds are also important as fattening 
food for wild boar. 



59 



'~~~— — ,.,__^^ Months 
Type of food""^ --^^..___^ 


1 


II 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


XII 


Reed (shoots) 


ЩШ 




^^ 


^ 
















m 


Reed (rhizomes) ^^^^^ 


л 


щт 






л 


^ 




^ 




^ 


Reedmaces (rhizomes) 






'Mi- 






^^ 


Smallreed (whole plants) 






X-. 










Giant fennel (roots) 












__.^ 






Millets (whole plants) 




















Saltwort (berries) 




















Herbage (greens) 


















Carp 














1 1 1 I 






^^и 






Locust 












1 












1 



Fig. 16. Seasonal changes in major food objects of wild boar in the lower course of 
the Hi River (from Sludskii, 1956). 

In the event of soil freezing these shoots become the main and often 
the only food throughout the winter. The green portions of herbage 
and grasses and racemes of cattail and other plants are of minor 
importance. 

Boar feed on a variety of food. Of the greatest importance is the 
Asiatic locust. In years of mass proliferation of this insect boar begin 
to feed on locust nymphs from mid-June; at the end of August they 
start collecting at places of locust concentrations, and later, until 
September, move in herds following locusts and consume large 
numbers of them when they lodge on the stalks of cane and other 
plants. Up to 1.5 kg of these insects have been found in the stomachs 
of wild boar. From October right up to the time of soil freezing, and 
sometimes until spring, wild boar continued to feed on dead locusts 
or dig up locust eggs. Italian locusts may also serve as an important 
autumn food (Sludskii, 1956). In northeast Cis-Caucasus (Kizlyar 
steppe and Caspian reeds) during the mass breeding of locusts in the 
1920's, wild boar almost completely changed over to feeding on these 
insects, and their feces consisted almost entirely of their remains 
(V.G. Heptner). 

In some cases wild boar consume large quantities of fish. In the 
Volga delta during April and May up to 1.5 kg of vobal [Caspian 
roach] and carp have been found in the stomachs of wild boar (Lav- 
rovskii, 1955). In several lakes and rivers of Kazakhstan wild boar 
catch carp in small shoals, open dried-out holes, and during winter 
freezes (Sludskii, 1956). In the Volga delta the animals regularly 
forage on fish, vagrant fledglings in colonies of herons and cormor- 



60 

ants and fledglings falling from nests. In small ilmens [Volga delta 
lakes] and kultuks [deeply indented shallow bays] wild boar gather 
bivalved mollusks, consume trapped muskrats, dig up burrows of 
mice as well as beetles and their larvae from the sand, destroy accessi- 
ble bird nests, and consume carrion. On the shallow northwest 
coasts of the Caspian Sea in the 1920's and 1930's wild boar regularly 
fed on mollusks at certain places, patiently wading through sea 
water throughout the night. At the Agrakhan spit, but mainly in 
Agrakhan Bay, a special method of hunting wild boar on moonlit 
nights has been developed, taking advantage of this behavior (V.G. 
Heptner). 
55 Living constantly close to water bodies and feeding on succulent 
diets wild boar always meet their water requirements satisfactorily. 
They visit water holes and traverse long distances to do so only 
during winter in the absence of snow. 

In some regions, for example the Caucasus, in the warm season 
of the year wild boar visit saline lakes (Nasimovich, 1938). Such 
lakes often serve as water holes for wild boar in which they gladly 
settle down to bathe and gather earthworms. The importance of 
saline lakes for wild boar and the regime of their visits to them have 
not yet been studied. 

Home range. Herds of wild boar, as well as single animals, are 
usually confined to permanent sites, the size of the herd depending 
on availability of food and shelters. Lone animals wander exten- 
sively over large areas of the habitat. Swine with small sucklings are 
confined to a small section. The size of the home range changes with 
the season. It is most extensive in autumn, especially in the event of 
nonavailability of normal foods, when wild boar visit farms. In such 
a case the animals will travel to plantations 6 to 12 km away (Din- 
nik, 1920; Cherkasov, 1867; Salikhbaev, 1950; Sludskii, 1956). 
According to Ivanenko (1956) wild boar daily visit feeding grounds 
located at distances of 16 to 18 km from their lairs, traversing every 
night distances of not less than 32 to 36 km in a straight line. This 
information appears dubious. In spring and summer the daily 
movement of herds of boar usually does not exceed 5 or 6 km as the 
crow flies. In winter, especially when snow is heavy, boar are con- 
fined to small sections of the home range. In Belovezh Forest the 
area of individual sections of home range of wild boar varies from 
0.5 to 2.5 km^ in winter; the animals daily traverse a few hundred 
meters to 3 to 4 km (Lebedeva, 1956). In the Far East the extent of 
daily travel depends on the availability of food and snow conditions 
and varies from a few hundred meters to 6 to 8 km (Rakov, 1956). In 
reed thickets along Hi River up to 100 animals were held up in an 



61 




Fig. 17. Furrows made by wild boar in forest clearance in Belovezh Forest. Photo- 
graph by L.S. Lebedeva. 

area 6.0 km x 3.6 km one winter because of heavy snow. In another 
case about 50 boar wintered in an area of 300 hectares, while 30 lived 
in an adjacent section of 120 to 150 hectares (Sludskii, 1956). Only 
during migration do wild boar traverse 20 to 40 km per day. 
56 Daily activity and behavior. In summer wild boar set out to feed 
before sunset and forage until dawn; males go out later than females 
with sucklings and young. High in the mountains boar are active 
during the day also. With the onset of cold, in October and 
November, the period of daily activity gradually shifts to daylight 
hours. In winter in most regions boar leave their lair in the morning 
and feed throughout the day; the lower the temperature, the later the 
departure of the animals for feeding. There are references to wild 
boar leading a nocturnal way of life even in winter in the southern 
part of the range. In spring, in April and May sows with sucklings 
leave the lair only during the day; at this time adults set out to feed 
in the evening, returning to the lair quite often at midnight and 
feeding for a second time early in the morning. Wild boar leave the 
lair invariably in the direction of the prevailing wind or across it 
and return against it. 



62 




Fig. 18. Spring lair of sows with sucklings and track leading to it. 
Belovezh Forest. Photograph by L.S. Lebedeva. 



Wild boar make their lairs in one section of the home range. In 
winter the structure of lairs is more complex than in summer when 
the animals quite often lie directly on the ground after raking up the 
soil or stones. In forests boar make lairs more often under the protec- 
tion of dense tops of trees or in dense undergrowth. Thus in the 
Caucasian preserve, of the 70 lairs examined, 40 were beside trunks 
of fir trees (Donaurov and Teplov, 1938); in Belovezh Forest, of 76 
lairs, 46 were under spruce and 13 in dense undergrowth (Lebedeva, 
1956). In Badkhyz wild boar like to lie under pistachio trees 
57 (Heptner, 1956). In mountains in winter the animals usually spend 



63 

their time on the sun-warmed slopes and in summer among large 
boulders or under rocks on the steep northern slopes. The winter lair 
is in the form of a depression among gathered dry twigs, moss, 
branches, pine needles, etc. In reed thickets the lair is in the form of a 
large heap of stalks and leaves; the animal either lies on top of the 
heap or burrows into it. The form and size of the lair varies. Males 
rest solitarily and their lairs have a thicker bed than those of females 
and sucklings. The elongated lair of the male corresponds to the 
body form of the animal using it. Lairs of females and sucklings are 
often circular. The female and her sucklings frequently rest in the 
same lair. Group lairs in which up to 10 to 15 animals spend the 
night are also known. In winter boar often rest under haystacks. The 
winter abodes of boar in eastern Siberia, Mongolia, Kazakhstan, and 
the Far East are more complex than those in Europe and the Cauca- 
sus. They are made up of large heaps of branches or reeds under 
cover of which the lair is built with a warm and dry bed of moss or 
dry twigs. The lair built prior to farrowing in which the litter 
spends most of at least the first two weeks, usually has a roof of 
branches of dry grass. Such lairs also have thick walls and a soft bed 
made of plant material. 

In area inhabited by boar there is always a place for wallowing in 
the form of a depression or pit filled with water and mud. Nearby 
soil is covered with tracks and adjacent trees, shrubs or rocks 
splashed with mud and scarred with animal scratchings. If there is a 
nonfrozen water body available, boar bathe in ail seasons of the year, 
but particularly so in the warm period when there are no rains, in 
the period of molt, and in the breeding season. 

Boar have relatively mobile middle toes and a distinct additional 
area of support in the form of the rear portion of the sole and hooves 
of the lateral toes. The weight load on the foot in adults" reaches up 
to 420 to 453 g (Nasimovich, 1955). Wild boar move relatively lightly 
along marshy, soft ground and shallow snow. However, the short 
limbs hinder movement in deep snow and the critical height of snow 
cover is 30 to 40 cm. When the height is more, boar, especially 
sucklings following their mother, try to walk in single file or use old 
tracks. At a snow height of over 70 cm adult animals move very 
slowly, making trenches for themselves (Nasimovich, 1955). Frozen 
snow crust is fatal to boar, injuring its legs. The wedge-shaped head 
and anterior body part enable the animal to move rapidly in dense 

"The weight load is 184 g per cm^ in 1.5-month-old sucklings and 375 g in 
juveniles six to seven months old. It is less among adult females than in males 
(Nasimovich, 1955). 



64 




58 Fig. 19. Muddy place for wallowing of wild boar in Belovezh Forest. Photograph by 

L.S. Lebedeva. 



reeds and shrub thickets by parting the stalks and branches. 

The olfactory sense is extremely well developed in swine and 
they can smell danger from a few hundred meters upwind; their 
hearing is also well developed, but their vision relatively weak. 

With the exception of adult males and females with small suck- 
lings, wild boar lead a gregarious life year-round; the young follow 
their mothers usually up to sexual maturity. The adult male tusker, 
outside the period of rut, is usually solitary and the name "loner" is 
appropriate. However, adult males have been known to stay with a 
herd of mothers and young ones during summer. 

There are five main factors which determine gregariousness and 
solitude year-round: 1) isolation of female before farrowing; 2) 
appearance of offspring and their separation during the course of 
the year; 3) adult males joining the herd during the breeding season; 
4) concentration of animals at places where fattening foods are avail- 
able; and 5) concentration of animals in the period of abundant 
snow. The latter two factors are not necessarily annual features and 
may be absent altogether in some regions. 

The herding instinct of wild boar is usually highest at the end of 
October or November, i.e., in the period of breeding when the males 



65 

join the herds of females and young ones. At this time the herd 
contains up to 5 to 8 aduk females, males and 15 to 20 young, 

58 making a total of 30 to 40 animals. On the average a herd at this time 
consists of 6 to 10 animals.^" At the end of mating, when the males 
separate from the herd, the herd size drops sharply. From January to 
March sows and young form groups of 3 to 5 each. In the event of 
heavy snowfall a concentration of animals occurs, especially toward 
the end of winter, when collective living facilitates their movement 
on snow and thus assists weaker animals, particularly the young, to 
secure food more rapidly. However, large herds are never formed 
since feeding on a small area would be difficult for a group of more 
than 8 to 12 animals. 

During the brief time of farrowing the herd size shrinks due to 
the isolation of sows with newborns. But by the end of April or May 
the herd size increases sharply, reaching in some cases the maximum 
for the year because of the appearance of the new brood. The pre- 
vious year's piglets again attach themselves to mothers along with 
two- or three-week-old sucklings; quite often two or three families 
merge into a single herd. From June to August families and young 
usually separate, thus reducing the herd number. At the end of 
August through September family herds and barren females may 
combine again at places where nut-bearing trees yield fruit and the 
herd thus increased once again. 

The annual herd number for Kazakhstan wild boar (1,127 ani- 
mals) averages 5.8 with a maximum of 7.7 and a minimum of 3.9 
(Sludskii, 1956). For the Caucasian preserve the corresponding 
values are 5.9, 10.2 and 3.3 (Donaurov andTeplov, 1938). For Belo- 
vezh Forest (17,340 animals) these values are 3.9, 5.2 and 3.0 respec- 
tively (Lebedeva, 1956). The mean size of the herd may vary 
perceptibly in different years and is determined by the total number 
of animals in the population, the size of the brood, variety and 
nature of the snow cover, and quantity of food. Thus in Belovezh 
Forest the average annual herd number over eight years of observa- 

59 tion varied from 2.2 to 5.2 (Lebedeva, 1956). For some months, espe- 
cially in winter and autumn (in subsequent years), instances of 
seven- to eightfold variations of mean herd size are known. 

Seasonal migrations and transgressions. Boar do not characteris- 
tically undertake regular seasonal migrations over most of their 



'''in the last century when hunting of wild boar in Central Asia was virtually 
non-existent, herds of boar numbering up to 50 to 100 animals were encountered in 
the breeding period in the lower course of the Syr-Darya (Skorobogatov, 1924; Sozon- 
t'ev, 1925). 



66 




60 Fig. 20. Herd of wild boar in Belovezh Forest. Photograph by V. Datskevich. 



range. In the plains of Europe, Kazakhstan, and Middle Asia wild 
boar often live year-round in the same region. However, some 
change of locality as well as concentration of animals at places 
where food is abundant probably occurs. The animals may gather in 
oak groves yielding acorns, leave river floodplains and go to salt 
marshes for fruits oiNitraria, or concentrate in the vicinity of culti- 
vated crops. At the same time, irregular migrations occur fairly fre- 
quently and are determined by the failure of the main food sources, 
high snow cover, soil freezing, and flooding or drying of lakes. In 
years of total failure of food sources, especially when the soil is 
covered with ice, migrations of wild boar are known from Belovezh 
Forest into adjacent regions; such phenomena are also known in 
Baltic region (Kartsev, 1903 etc., Ling, 1955). 

Movement of wild boar from the Volga delta to the Prikumsk 
valleys downstream, Volzhsk-Ural sands, and Kamysh-Saman lakes 
have been described (Karelin, 1875; Rukovskii, 1947). Winter migra- 
tions are a frequent phenomenon between the Syr-Darya and Amu- 
Darya when the animals go to the sands of Kyzylkum during intense 
freezing of the muddy soils of reed clumps. Similar instances of 
movement of wild boar to a distance of up to 100 to 200 km are 
known on the Chuya River from where the animals go to Muyun- 
kum. Hi River, and several other regions (Bazhanov, 1945; Shul'pin, 
1948; Sludskii, 1956; and others). 

In most montane regions wild boar perform regular seasonal 
vertical migrations associated with seasonal availability of food and 



67 

nature of snow cover. Wintering wild boar utilize the middle and 
lower regions of hills but move to the foothills in winters of heavy 
snow. Boar descend to the middle and lower regions even in autumn 
when seasonal foods, mostly nuts and fruits, are abundant. In the 
event of unusual snowfalls in midwinter additional migrations into 
regions with less snow take place. In April and May a reverse migra- 
tion occurs. The range of vertical movements varies from a few 
hundred meters to 1,000 to 1,500 m in the Caucasus, 2,000 m in the 
Tien Shan, and reaches 60 to 80 km in Sikhote-Alin (Dinnik, 1910; 
Nasimovich, 1936, 1939 and 1955; Zverev and Keleinikov, 1947; 
Shul'pin, 1948; Salmin andShamykin, 1949; Abramov, 1954; Rakov, 
1956; Chernyavskaya, 1956; and others). 

Reproduction. In most regions mating of wild boar occurs from 
November to January, but the duration of intense rut in a given year 
and place usually does not exceed one to one-and-a-half months. In 
males the commencement of rut is preceded by the development of a 
"stuffing" ("fat"), in the form of a connective tissue layer of 2 to 3 
cm thickness under the skin on both sides of the shoulder blades up 
to the last rib. These "pads," which begin to grow roughly two 
months before rut, protect the male from deep wounds which could 
be inflicted by the tusks of rivals during combat. At the commence- 
ment of rut the testes enlarge to almost double their normal size and 
the glands secrete a viscous, foamy, yellowish liquid, often sprayed 
on tracks. A male in search of a herd with females in estrus wanders 
widely and feeds little. On encountering a herd it drives away young 
piglets and sucklings and starts chasing a female in estrus, often 
pursuing her in circles. A male on finding a herd with females will 
vigorously fight any subsequent competitor. At the end of rut males 
are often severely mauled and emaciated. During rut they lose about 
20% of their weight (Markov, 1932). Each adult male generally gath- 
ers one to three females, indulging in limited polygamy. The 
number of females to a male may reach 5 to 8 and, in the period of 
rut, the herd may number 30 to 40 animals, including sucklings and 
young pigs (Sludskii, 1956). 
60 The period of rut varies insignificantly in relation to the geogra- 
phic position of the region. Usually rut sets in first in western Euro- 
pean animals, at the end of October or early November, and last in 
Trans-Baikal, at the end of November or in December. Young ani- 
mals come into rut two or three weeks after older ones. The period of 
rut may shift sharply to a much later period with a failure in main 
sources of food. In Belovezh Forest when total failure of food supply 
occurred, rut was observed only in February and March instead of 
November and December (Lebedeva, 1956). A delay of one month in 



68 

rut in wild boar was recorded around the Hi River as a result of 
scarcity of food (Sludskii, 1956). Instances have been described 
(Oloff, 1951) when the mating of wild boar occurred in western 
Europe in March and April and once even in July and August 
because of the nonavailability of food.^^ 

There are references in literature to extremely well-fed boar also 
coming into rut late, but more so domestic swine. This pheno- 
menon is probably associated in warm winters in the Caucasus with 
abundant food supply (Loginov, 1936; Sludskii, 1956). In some cases 
rut evidently sets in twice, as evidenced by the presence of very small 
sucklings in January and again in April (Kissling, 1925; Oloff, 1951; 
Lebedeva, 1956). 

Females usually come into estrus for the first time in the second 
year of life, at the age of 18 to 20 months; males come into rut in the 
fourth or fifth year of life. However, physiologically females attain 
sexual maturity in the first year (8 to 10 months) and males in the 
second year.^^ Males before the age of four or five years are not 
permitted to mate by the more powerful tuskers. In western Europe 
only in years of very good availability of food up to 50% of young 
females take part in reproduction. In years of moderate food supply 
the corresponding figure is 10%, and in years of poor food supply 
young females produce no offspring (Oloff, 1951). In Tadzhikistan 
corpora lutea were detected in some young females, indicating not 
only their sexual maturity but also their impregnation at the age of 8 
to 10 months (Ivanenko, 1956). However, the embryos were usually 
resorbed as the females at that age were poorly fed. The participa- 
tion of young females in reproduction under favorable conditions of 
food supply is also known in other regions. 
61 The duration of gestation among wild boar has not been accu- 
rately established. In old animals it is 133 to 140 days and in those 
reproducing for the first time 114 to 130 days (Kissling, 1925; Ken- 
neth, 1943). Mass farrowing occurs from March to May, more often 
in April. In Europe young usually emerge in March to early April 
and in Trans-Baikal at the end of April to early May. Corresponding 
to the change in periods of mating (see above), periods of birth of 
young also change, being determined by the age of the mother and 
availability of food. 

''No doubt the situation aicountered by Flerov and Gromov (1935) who record 
February and March as the period of mating and correspondingly May and June as 
the period of farrow. 

'^Ivanenko (1956), on the basis of a notable enlargement of testes and prostate 
gland, asserts that males also attain maturity by November or December in the year of 
their birth (8 to 10 months). 



69 

Litter size also depends on the extent to which the female has fed 
and her age. The litter size of young females is roughly half that of 
adult females. Depending on the availability of the main sources of 
food, the number of young in a litter may vary from 2 to 10 or more. 
On the average swine produce 4 to 6 young. The average number of 
young in a brood over a 50-year period for western Europe was 5.5 
(Oloff, 1951); in Belovezh Forest for 8 years — 5.9 (Lebedeva, 1956); 
Caucasian preserve — 4.4 (Donaurov and Teplov, 1938); Far East — 
6.4 (Salmin and Shamykin, 1949); Volga delta — 6.0 (Lavrovskii, 
1955); Kazakhstan— 5.4 (Sludskii, 1956); northern Mongolia— 4.0 
(Bannikov, 1954); and Tadzhikistan — 3.9 (Ivanenko, 1956). The 
maximum known number of young in a brood is 10 to 12. Instances 
have been recorded of a mother with 14 to 16 sucklings but it is 
highly probable that the offspring of another, dead sow simply 
attached themselves to a new mother. 

The number of fertilized eggs significantly exceeds the number 
of offspring. In Tadzhikistan 4 to 15 corpora lutea were found in 6 
adult females (average 9.6), 2 to 9 embryos in 12 pregnant sows 
(average 4.3), and 2 to 10 offspring born to 8 females (average 3.9) 
(Ivanenko, 1956). Hence a large number of embryos are resorbed. 
Evidently in the years of failure of food sources the number of 
resorbed embryos increases, which also determines the size of the 
litter. In western Europe in the years of nonavailability of food the 
number of young in a brood has averaged 3.0, reaching 8 or 9 in 
years of plentiful food, with along-time mean of 5.5 (Oloff, 1951). In 
Belovezh Forest in years of poor availability of food the average 
number in a brood has been 3 or 4, reaching 8 to 10 in years of 
plentiful food, with a long-time mean of 5.9 (Lebedeva, 1956). Such 
phenomena have also been recorded for the Caucasus, Far East, 
Kazakhstan, and other regions. 

The number of barren females varies even more but precise data 
are scanty. In Tadzhikistan 5 of 13 dissected females, i.e., about 40%, 
were barren (Ivanenko, 1956). In the Caucasian preserve on the aver- 
age 50% were barren (Donaurov and Teplov, 1938). In western 
Europe over a 50-year period the ratio of fertile to barren females 
averaged 3 : 7, i.e., about 60% were barren. In years of abundant 
supply of food the number of barren females dropped to 25%, while 
in years with poor food supply it rose to 80% (Oloff, 1951). An 
increase in percentage of barren females in years of scarcity of food 
has also been recorded in Kazakhstan, the Far East, and other 
regions. 

Growth, development, and molt. In the first week of birth the 
young do not leave the lair and, in the absenceof their mother, lie 



70 

closely pressed to each other. Quite often the mother partly covers 
the litter with bed material before she leaves the lair. Commencing 
from the second week the young move with the mother, returning to 
the lair for rest. The young reside in the same lair with the mother 
up to the age of one year, sometimes more, especially in cold winter. 
From about the age of two weeks sucklings run away and quickly 
hide in the grass or stand immobile in dense shrubs or reed thickets 
when threatened with danger. The stripes on their body, persisting 
up to July, camouflage them well. 

The lactation period lasts from 2.5 to 3.5 months. However, at 
the age of two or three weeks the young begin to dig up soil, gather- 
ing small bits of rootstock and small soil animals. They attempt to 
dig up soft soil even at the age of one week but evidently do not 
succeed in gathering food. A small amount of tubers and plant 
material has been found in the stomachs of two- week-old piglets. 

Newborn piglets weigh 600 to 1,000 g; the average weight is 750 
62 g. In the lower courses of the Hi River in May sucklings weigh 2.0 to 
4.5 kg, in June — 3.7 to 9.5 kg, in August — 19.5 kg, and in December- 
January — 16 to 30 kg, with an average of 22.5 kg. This is followed by 
a small drop in weight in February, finally rising to 40 kg toward 
the end of the first year. The weight of adult females is 65 to 100 kg, 
average 83 kg, and of males 95 to 200 kg, average 126 kg (Sludskii, 
1956). In Tadzhikistan sucklings in summer weigh an average of 
16.4 to 17.1 kg, in autumn 31 to 33 kg, and in winter 35 to 38.2 kg. 
Young pigs weigh 38 to 44 kg. Adult females weigh 71 to 123 kg and 
males 74 to 144 kg (Ivanenko, 1956). The weight of boar in tugais of 
Tadzhikistan does not exceed 100 to 120 kg; in the mountains some 
cases are known of animals weighing 158 kg (Chernysev, 1958). In 
northern Mongolia (Khentei) the weight of sucklings is 20 to 25 kg 
by September and of young by spring 27 to 33 kg (Bannikov, 1954). 
In central Europe the average weight of young pigs of different ages 
based on long-term data was 19.3 to 28.4 kg, of juveniles 35.9 to 42.8 
kg, of adult females 44.6 to 61.4 kg, and of males 54 to 77 kg. The 
weight of wild boar of all ages varies sharply in different years 
depending on the availability of food. The maximum weight of boar 
for different regions in central Europe was 153 to 230 kg (Mager, 
1941; Oloff, 1951; and others). In the seventeenth century captured 
wild boar reportedly weighed 225 to 230 kg (Brandt, 1912). In Latvia 
the largest animals weigh up to 200 kg but one tusker weighed in at 
236 kg (Kalnin'sh, 1950). In Belorussia a male weighing 246 kg was 
killed in 1951 (Serzhanin, 1955). In the Volga delta in the last cen- 
tury wild boar weighed 48 to 112 kg with a maximum of 190 kg 
(Yablonskii, 1875). In other regions of the Caspian coast they tipped 



71 

the scales at 300 to 320 kg (Karelin, 1875). At present the maximum 
weight of boars in preserve is 250 to 270 kg (Lavrovskii, 1955). 

In Middle Asia (Syr-Darya) in the nineteenth century the maxi- 
mum weight of males was 128 to 160 kg and very rarely 190 kg 
(Severtsov, 1874). In the Caucasian preserve the weight of males was 
64 to 178 kg with an average of 166 kg; females weighed 48 to 103 kg 
with an average of 68 kg (Donaurov and Teplov, 1938). At the end of 
the nineteenth and early in the twentieth century (Dinnik, 1910) 
males attained weights of 240 to 255 kg and females 120 to 145 kg. In 
Azerbaidzhan the weight of adult males reaches 142 kg with an 
average of 114 kg and for young male pigs 68.5 and 63 kg respec- 
tively; for adult females these values are 98.5 and 86.5 kg, and for 
young female pigs 65.6 and 58.5 kg (Safarov, 1958). In eastern Sibe- 
ria in the last century boar reached 240 to 270 kg (Cherkasov, 1867). 
In the Far East large boar weighed 150 to 170 kg, some animals 
weighing up to 300 to 320 kg (Abramov, 1954). 

Newborn piglets have one milk incisor and one canine on each 
half of the jaw. At the age of 2.0 to 2.5 months two premolars appear 
on each half of the jaw. By autumn, i.e., age six months, all the 
incisors (3) and premolars (4) have formed. By winter the milk 
canines and three milk incisors are replaced by permanent teeth; the 
first molars have also appeared. Yearlings and two-year-olds have 
only permanent teeth and start developing the second molar. The 
third (last) molar is seen at the age of two to three years. In the 
middle of the fourth year all the teeth, except the canines in males, 
cease to grow, while the first and second molars already have worn- 
out crowns (Ivanenko, 1955). The growth of canines does not cease 
in old males and they become very powerful, increasingly curved, 
acquiring a semicircular form with advancing age. 

The life span of wild boar in nature is 10 to 12 years. In western 
Europe, however, animals older than 6 to 8 years are found very 
rarely. In Tadzhikistan of the 207 living boar, those older than 6 
years constitute only 4.3% (Ivanenko, 1956). References to wild boar 
living up to 20 years refer to animals living in captivity. 

Pigs are born with only bristles; the underfur forms by the age of 
two weeks. The growth of underfur and the wearing down of ends of 
embryonic hair (bristles) causes the disappearance of body stripes by 
the time the piglet is three months old. The maximum growth of 
hair cover is observed in eight-month-old sucklings. 

Wild boar molt for the first time one year after birth, i.e., between 
63 March and May.^'' Molt commences with the shedding of underfur; 

''The statement by Sludskii (1956) that young boar molt for the first time at the 
age of 3.5 to 4.0 months is erroneous. 



72 

bristles fall thereafter. New bristles form immediately while the 
underfur develops only toward the end of July or in August. During 
June and July only the crown of bristles develops, which explains 
the silvery-white summer coloration of the animals. By September 
the hair cover has fully developed (Ivanenko, 1956). 

Molt in adult boar proceeds in the same manner as that in young. 
Well-fed males molt first, followed somewhat later by emaciated 
animals and lactating females. Molt in the latter may be delayed up 
to early June. Compared to wild boar of Europe and Middle Asia in 
eastern Siberia and the mountains molt is somewhat delayed until 
spring. 

Enemies, diseases, parasites, mortality, competitors, and popula- 
tion dynamics. Wolves are the main enemy of wild boar over much 
of its geographic range. In Belovezh Forest, in spite of its rich ungu- 
late fauna, wild boar are preyed on by wolf more often than any 
other animal. Remains of boar were found in 24.9% of all the data 
(1,063) on food of wolves; in various years the percentage ranged 
from 7.5 to 39.5. In winters of abundant snow wolves inflict the 
greatest damage on boar (Gavrin and Donaurov, 1954). In the Baltic 
region the boar population largely depends on the population of 
wolves, and in severe winters the- predators eliminate their prey 
almost completely (Kalnin'sh, 1950; Ling, 1955). In the Caucasian 
preserve boar was found in 37.6% stoinachs of wolves (154). In win- 
ters of heavy snowfall wolves quite often destroy herds of pigs, 
including sucklings and young pigs (Donaurov and Teplov, 1938). 
In Kazakhstan (lower courses of the Hi River) boar remains were 
recorded in 22.3% of all available data (1,052) on food of wolves. One 
adult wolf can destroy between 50 and 80 boar of different ages in 
one year (Sludskii, 1956). The significance of wolf as an enemy of 
wild boar is high in other regions also. Wolves kill mainly sucklings 
and young pigs, rarely attack sows, and avoid attacking tuskers. 
Instances are known of the death of predators as a result of injuries 
inflicted by the tusks of swine. Dense shrubs and cane thickets offer 
good hiding places for wild boar and hence wolves very rarely attack 
tuskers or sows. 

All the remaining predators are of lesser importance in the de- 
struction of wild boar. In several regions lynx attack piglets and 
sucklings. In Belovezh Forest boar remains were found in 8.3% sto- 
machs (of 23 examined) and in 3.9% feces (of 206 examined) of lynx 
(V.F. Gavrin). Instances are known of attacks by lynx on young boar 
in the Caucasus, Far East, and eastern Siberia (Dinnik, 1914; 
Abramov, 1954; Cherkasov, 1867). In rare instances sucklings are 



73 




64 Fig. 21. Wild boar tracks in mud in reed dumps on Chuya River. 

Photograph by A.G. Bannikov. 

attacked by jungle cats and spotted cats* (Sludskii, 1956). Instances 
are known of attack of sucklings by snow leopard (Shul'pin, 1948) 
and leopard (Dinnik, 1914; Bil'kevich, 1924; Baikov, 1927). In the 
last century in Trans-Caucasus, Kazakhstan, Middle Asia, and the 
Far East wild boar constituted the main quarry of tiger; however, at 
present this predator has either disappeared or become extremely 
rare and hence its role in the destruction of wild boar is minimal. In 
the Far East brown bear and yellow- throated marten sometimes 
attack piglets and sucklings (Abramov, 1954). Evidently bear and fox 
feed on the carcasses of dead animals; it is possible that bear attacks 
on living animals are exceptional. Instances of killing of sucklings 
by stray dogs are reported quite often. 

•Species referred to by this common name is unclear — Sci. Ed. 



74 

About 20 species of parasitic worms have been identified for wild 
boar (Badanin, 1931; Pavlovskii, 1946; Dubinin, 1952; Rukhlyadev, 
1952). Maximum infection is observed in summer. The most danger- 
ous helminths are Metastrongylus parasitizing the lungs and caus- 
ing mortality among sucklings. Metastrongylus infections occur 
through an intermediate host, the earthworm. Of pathological 
importance to wild boar is the fluke Gasterodiscoides hominis, 
which also parasitizes man. Wild boar are likewise known to harbor 
Trichinella spiralis and instances have been described of infection of 
man by this parasite through the consumption of meat of wild boar 
(Chun-Syun, 1958). Taenia solium and Balantidium coli are also 
known to parasitize wild boar (Appasov, 1958). 

Ticks (Dermacentor, Rhipicephalus, and Hyalomma) and a spe- 
64 cies of hog louse {Haematopinus sus) parasitize wild boar quite 
often, especially in the southern regions (Pomerantsev, 1946; Pav- 
lovskii, 1946). The animals also suffer acutely from blood-sucking 
Diptera. To escape dipteran attacks wild boar spend considerable 
time in water, move from reed beds into the steppes, hide in dense 
shrubs, and so on. 

Swine plague spreads faster than any other disease of wild boar 
and causes extensive mortality. Epizootic diseases are well known in 
Germany, Poland, Hungary, Belorussia, the Caucasus, the Far East, 
Kazakhstan, and other regions (Kissling, 1925; Kartsev, 1903; 
Donaurov and Teplov, 1938; Smirnov, 1928; Abramov, 1954; 
Kovach, 1957; Mom berg, 1954; Sludskii, 1956; and others). The 
extremely contagious foot-and-mouth disease, known among wild 
boar of western Europe, the Caucasus, and Kazakhstan can assume 
an extensive epidemic character (Dinnik, 1910; Gutira and Marek, 
1931; Sludskii, 1956). Pasteurellosis or hemorrhagic septicemia 
(Smirnov, 1928) and tularemia are rare; wild boar contract tularemia 
either from rodents or through blood-sucking insects (Pashova, 
1950). Siberian ulcers are also very rare (Andreev and Andreev, 1948). 
Swine erysipelas is contracted by wild boar from rodents or through 
hog lice or ticks (Smirnov, 1928). Wild boar also harbor pyroplasmo- 
sis and Aujezsky's disease (Skomorokhov, 1951; Andreev, 1948). In 
Czechoslovakia in 1950; and in Poland in 1954, wild boar suffered 
from epidemic viral teschen disease (porcine enteroviral disease) 
(Shaflyarskii, 1955). 

In years of abundant availability of food wild boar have no 
serious competitors in much of their geographic range. In years of 
poor availability of acorns in mixed forests in the northern part of 
the range a large number of animals such as deer, squirrels, jays, etc. 
compete with boar for acorns. In eastern Siberia a similar pheno- 



75 

menon occurs with regard to pinenuts. In the Caucasus squirrels, 
after their acclimatization, began destroying beechnuts which had 
not yet fallen. By consuming an appreciable part of the crop they 
became an acute competitor of wild boar. 

Twice within a decade a large number of wild boar died in the 
Volga delta during floods when the animals became emaciated for 
want of adequate food and remained in water for many days (Dyu- 
nin, 1926; Isakov, 1951; and others). The young particularly die in 
large numbers in swollen waters due to spring — summer floods in 
this region, causing a sharp reduction in wild boar population (Lav- 
rovskii, 1955). Such phenomena have been observed in the lower 
course of the Syr-Darya River (Sludskii, 1956) and rarely in other 
rivers. 

Fires lead to the death of wild boar in Middle Asia, Kazakhstan, 
and along the Caspian coast. During fires in cane and tugai thickets, 
especially with strong winds, a significant number of sucklings per- 
ish. Furthermore, fires deprive the animals of shelters, making them 
more vulnerable to wolves (Sludskii, 1956). 
65 The population dynamics of wild boar are characterized by sharp 
numerical changes over short intervals of time. Inadequate food 
supply combined with deep snow or ice-crusted ground exert a per- 
manent influence on population attrition. The activity of wolves is 
important everywhere. Years of epidemics and natural calamities 
(floods and fires) often lead to the death of an appreciable percentage 
of local populations. Most often mortality of wild boar is the result 
of a combination of unfavorable factors. For example, in the Polish 
part of Belovezh Forest over 60% wild boar perished in the winter of 
1955 to 1956; the surviving swine were so emaciated that they could 
not breed in spring. The causes of death were soil freezing, forma- 
tion of three-layered frozen snow crust, and a high snow cover, all of 
which prevented the animals from getting at their food. Moreover, 
rodents depleted the small yield of acorns. In this winter swine ery- 
sipelas, which usually affects them lightly, led to the death of wild 
boar already weakened by inadequate food (Tsabon', 1958). 

A rapid increase in population is assured by the high fertility 
potential of wild boar, large-sized litters, and the early onset of sex- 
ual maturity. However, litter size, the age at which females repro- 
duce for the first time, and the percentage of females participating in 
reproduction vary sharply, depending on the state of the food base 
(see above). A population of wild boar can dwindle to one-tenth or 
double or treble in just a few years. 

Precise data on the mortality of young swine are scanty. In Belo- 
vezh Forest an average of 64% survive to their first birthday (Lebed- 



76 





., ~^f... 



Fig. 22. Tracks of wild boar in snow in Belovezh Forest. Photograph by 
A.G. Bannikov. 



eva, 1956); this figure in the lower course of the Hi is 60% (Sludskii, 
1956), and in Khentei about 40% (Bannikov, 1954). When food is 
abundantly and readily available mortality of young is low and over 
90% survive up to the age of sucklings. Contrarily, in years of inade- 
quate food supply, especially when combined with heavy snows and 
soil freezing, not more than 20% survive to spring. Evidently every- 
where the maximum death of sucklings occurs in late autumn and 
early winter and is determined by worsening conditions of food 
accessibility, especially during intense frosts. In summer, when suck- 
lings take over feeding themselves, become more mobile, and often 



77 

run about independently, a second peak of mortality has been 
recorded (Danaurov and Teplov, 1938; Lebedeva, 1956). 

Among adult wild boar the sex ratio is close to 1:1 in most 
regions. However, among newborns females predominate and con- 
stitute 60 to 70% of total births. This reveals a high mortality of 
females at an early age. 
66 The age structure of the herd changes perceptibly in different 
years. Usually, the year's brood form 50 to 60% and sucklings 9 to 
12%. Following severe winters, when sucklings are the first to die 
and females give birth to small-sized litters, the percentage of piglets 
and sucklings decreases three or four times compared to the average, 
and hence piglets and sucklings together form only 5 to 10% of the 
herd. Contrarily, after a favorable winter the herd acquires a percep- 
tibly youthful appearance. 

Field characteristics. Coloration is extremely variable but from a 
distance wild boar often appear almost black; in summer the bristles 
gleam in the sun and light, sandy-colored specimens are seen. 

Hoofprints of adult wild boar in soil invariably show the 
imprint of the lateral digits; in small sucklings traces of only two 
digits are left. The print length of an adult boar ranges from 16 to 25 
cm, generally about 20 cm; the width is even more variable but more 
often about 18 cm (maximum distance between lateral digits). The 
winter prints are deep. The animals drag their legs and often plough 
a continuous furrow in the snow. Lone males on selecting a site for 
lying up turn around, executing a semicircle, and lie down with 
their head in the direction of the track at the point of entry; females 
do not turn around in such a manner but lie down straight on the 
track. Wild boar often lie down at the foot of trunks of densely 
leaved trees, below upturned trees, in dense undergrowths, shrubs, 
and in breaks in cane groves after making a suitable bed (see above). 
The animals make deep tracks leading to places of feeding and wal- 
lowing (see above). The habitat of wild boar is characterized by deep 
furrows in the soil of meadows, forests, or cane thickets, ploughed to 
a depth of 10 to 20 cm. The feces of wild boar in autumn, winter, and 
most of the summer are in the form of three or four flat cakes, 2.0 to 
2.5 cm in diameter, joined together; of these one is larger (thicker) 
than the rest. In spring and early summer feces often lack definite 
form. 

While resting adult males are silent, females grunt often, and 
piglets whine. A frightened boar makes a loud noise ("ukh! ukh!") 
or emits a screech ("gu-gu-gu!"). While feeding boar wave their tail 
rhythmically. On sensing danger the tail hangs down or rises in the 
form of a question mark, the animal stops feeding, and raising its 



78 




Fig. 23. Tracks of female wild boar with brood in Belovezh Forest. Photograph by 

L.S. Lebedeva. 



snout gulps air which gives rise to an audible puffing sound. (A.B.) 

Economic Importance 

The extent of catch of wild boar in the past and at present in the 
Soviet Union is difficult to assess since the main part of tjie catch is 
retained by hunters. In pre-Revolutionary Russia about 50,000 
animals were caught, mainly in the Caucasus, along the Caspian 
coast, and in the southern part of the Far East. In Middle Asia wild 
67 boar was caught less as its meat was not used for food; rather it was 
killed to protect melon fields and plantations. 

At present about 7,000 to 8,000 swine are caught annually in the 
Caucasus (Vereshchagin, 1947), 6,000 to 7,000 in Kazakhstan (Sluds- 



79 

kii, 1956), and about 5,000 in Middle Asia. No information is availa- 
ble for other regions but evidently the total for the country as a 
whole is currently between 60,000 and 70,000 per year. Data per unit 
of land are not available. In Germany at the end of the nineteenth 
century 0.08 to 0.1 animals were caught per 100 hectares. At the end 
of the 1930's this figure had risen to 0.21 to 0.26. Between 1935 and 
1939 a total of 33,000 animals were caught (Oloff, 1951). 

In the past the main methods of catching were extensive "beats" 
and hunting with dogs. Beats were done on foot as well as on horse- 
back. Beaters moved in a chain or rode through the forest or cane 
thicket driving the animals toward a group of hunters. In the desert- 
steppe regions thickets were often set on fire. Setting fires for the 
purpose of hunting is now prohibited. During the hunt the animals 
were killed with lances and spears; shooting came later. 

Hunting with dogs, when a large pack of up to 20 or more strong 
freely scoured cane thickets and forests, was extensively practiced 
earlier; now dog packs are prohibited. In earlier days a large number 
of piglets and sucklings were frightened by the dogs and killed. In 
hunting with a pair of dogs, they are set only on the trail of an adult 
animal. The task of the dogs is to stop the boar and hold it at bay 
until the hunter arrives. Also prohibited now are methods using 
traps, loops, crossbows, and pit traps. At present hunting on the sly 
is commonly practiced everywhere. The hunter follows a fresh track 
and, orienting himself by sounds, approaches the boar against the 
wind. It is extremely difficult to approach a silent and wary solitary 
male. Hunting by lying in wait is also practiced equally extensively. 
At the feeding grounds of wild boar, often in plantations, groves, 
and other places, pits are dug, hides set up, scaffolds constructed on 
trees, etc. where the hunter waits for the animal, taking advantage of 
the prevailing wind direction. This type of hunting is sometimes 
combined with beats. 

At present the period of hunting in most regions extends from 
September I through December 1. The period from September 15 
through November 15, i.e., after the cessation of lactation and before 
some of the females become pregnant again, is a more correct time. 

Meat, hide, and bristles are the products obtained from wild 
boar. The weight of meat in summer amounts to 55% and in winter 
65% of body weight. According to meat-dressing organizations the 
body of a boar should yield an average of 50 kg meat. A large boar 
may yield 15 to 20 kg fat and, exceptionally, even up to 30 kg or 
more. The hide of a large boar can stretch up to 300 dm^; it yields 350 
to 1,000 g bristles and about 400 g of "fuzz" 

When wild boar live in the neighborhood of townships, gardens. 



80 

and farms, they cause damage by digging up potatoes, damaging 
melons, watermelons, and maize, eating the crop from the stage of 
milk formation onwards. In the latter case damage is due mainly to 
trampling considerable areas in plantations (Kartsev, 1903; Satunin, 
1906; ShilHnger, 1928; Markov, 1934; Kalnin'sh, 1950; Abramov, 
1954; Sludskii, 1956; Kissling, 1925; Snethlage, 1949; and others). 
However, the damage inflicted by wild boar is often exaggerated 
(Kissling, 1925; Ling, 1955). Only in small sections of farms situated 
among forests or cane thickets, when the wild boar population is 
high, do the animals inflict significant damage on cultivated plants. 
The extent of availability of natural foods is of great importance; 
when natural food is not available, boar enter farms in large 
numbers and damage is severe. In Belovezh Forest in years of moder- 
ate availability of natural foods 34 to 47% of the population enter 
farms. In clearings animals damage potato and buckwheat crops up 
to 30%, but on the whole the damage to fields is 0.08 to 4.7% (Lebed- 
eva, 1956). To protect farms from wild boar, in Central Europe 
several measures have been suggested, including distraction and 
fright (Snethlage, 1949). In Kazakhstan leashing dogs to long chains 
at places where boar enter the plantation has been suggested as the 
most effective method of protection (Sludskii, 1956). 
68 Wild boar play a significant role in afforestation but this aspect 
has not been studied adequately. The number of seeds which a swine 
consumes in years of moderate availability does not delay seeding of 
a forest and the damage inflicted by wild boar in this process is 
imperceptible (Oloff, 1951). The digging activity of boar is very 
important to the life of a forest. By digging up large areas the ani- 
mals promote the sowing of seeds which, in turn, leads to restoration 
of wood-forming species. The role played by wild boar is very 
important therefore in the case of mossy fir stands of the Tien Shan 
(Korelov, 1947), nutpine groves of eastern Siberia (Abramov, 1954), 
and oak forests of Europe (Snethlage, 1949; Lebedeva, 1956). The 
extremely positive role of wild boar in afforestation is evident when 
their population is not high. However, in the event of too high a 
density they plough up the same section over and over and dig up the 
germinadng seeds. They also promote a shift in wood-forming species, 
as observed in Central Europe where oak forests were transformed 
into birch due to their activity (Fitingof-Rish, 1952). In Belovezh 
Forest wild boar promote the replacement of oak by spruce (Lebedeva, 
1956). 

The importance of wild boar in the destruction of forest pests has 
been well known in western Europe for a very long time (Hennert, 
1797; Spangenberg, 1818; Ratzburg, 1856; Fohlisch, 1880; Mei- 



81 

maiter, 1915; Gaber, 1950; Klemm, 1951; and others). The impor- 
tance of wild boar is particularly notable in the destruction of larvae 
of June bugs and Cephalis otitis; the animals reduce these popula- 
tions by 20 to 30% (Fitingof-Rish, 1952). Boar also destroy the pine 
geometrid moth and other forest enemies. When there are local 
spurts in population of forest pests, wild boar suppress them and 
prevent their outbreaks. However, if an outbreak has already 
occurred in a large area, wild boar are unable to restrict its further 
spread (Tsappe, 1958). There are no concrete data to demonstrate the 
importance of wild boar in the destruction of forest enemies within 
the Soviet Union. 

At the end of the last and the commencement of the present 
century small groups of wild boar were released time and again in 
private farms around Moscow. Experiments with acclimatization 
were carried out in 1937 in Zavidov hunting area, north of Moscow, 
by releasing nine boars. In 1940 there were about 100 of them and by 
1947 their number exceeded 300. The animals had dispersed widely 
and were found at places tens of kilometers away from their place of 
release. Hence only some of them visited the original feeding points 
(I. Lyapunov). Between 1948 and 1949 in the hunting zone of Rya- 
zan' district 40 boar were released. They prospered and dispersed 
into neighboring districts; some animals moved 60 to 180 km from 
the point of release but evidently all of these died. Wild boar were 
released in small numbers in the 1940's and 1950's in other farms of 
Moscow, Yaroslav, and adjoining districts. In 1957 a total of 35 Far 
East wild boar were released in the Crimean hunting preserve 
(Tkachenko, 1959). In the warm season wild boar find adequate nat- 
ural food everywhere, but in winter are compelled to supplement this 
food, the extent of which is determined by the amount of natural 
foods available and their accessibility (height of snow cover and soil 
freezing). Even when food supplements to boar were stopped, the 
animals were still found six years later but subsequently disap- 
peared. In addition to supplying food supplements, extensive protec- 
tion of wild boar farms from wolves is essential. The permissible 
density of wild boar in mixed forests in European sectors is 6 to 12 
animals per 1,000 hectares. In farms where muskrats are present, 
wild boar destroy their burrows and feeding places in winters of ice 
crusts and hollow ice (Pokrovskii, 1950; Novikov, 1957). 

In captivity wild boar get on well but young sucklings without a 
mother grow slowly and poorly. Wild boar interbreed readily with 
domestic swine; in nature interbreeding is known in the Caucasus, 
Belovezh Forest, the Far East, and other regions. Hybridization 
experiments with large white swine and wild boar in Kazakhstan 



82 

established that the resukant hybrids possess high vitahty. Hybrids 
of the third generation feed on less concentrated types of food, record 
very high (by 14 to 15%) weight increase, and consume less food per 
kg weight increase. Their skeletons are fattier than those of the large 
white variety. Hybrid swine adapt well to the utilization of summer 
pastures in southeast Kazakhstan (Butarin, Es'kov, Miletskii and 
Bagrovskaya, 1955). (A.B.) 



Suborder TYLOPODA Illiger, 1811 



69 Tylopoda are artiodactyls of large to moderate size with relatively 
elongated limbs and either of heavy or light build.' 

These animals walk on the two toes present on each leg. The 
lateral toes are totally reduced and the metacarpal or metatarsal 
elements totally absent. The remaining corresponding bones of the 
toes fuse into a cannon bone, which is bifurcate at the end and has 
no paired blocklike articulating surface. Terminal phalanges small 
and rounded. Hooves absent, replaced with only small, blunt, 
curved nails. Undersurface of foot consists of a broad, paired or 
unpaired, elastic, callused, cushionlike padding. Ulna and radius 
fuse in the proximal and, in some species, even in the distal part; 
fibula partly reduced. 

Skull with an extremely elongated frontal portion. Orbit closed 
from behind. Ethmoid pit well-developed. Tympanic bulla with 
lamellar osteosepta. Sagittal crest present. Horns absent and mastoid 
bone exposed. Condyle of lower jaw spherical. 

Young camels have three upper incisors on each side, while 
adults have only one canine-like upper incisor on each side. Lower 
incisors total three, the third in some animals also resembling a 
canine. Canines typical in form (conical), present in both jaws, and 
separated by a diastema from premolars. Lower canine separated 
from incisors. First premolar in upper jaw has the form of a canine in 
some camels and is separated by a diastema from molars. Latter with 
crescent-shaped cusps (selenodont) and hypsodont. Dental formula: 

i I- , с ]- , pm '^^ , m I = 34 to 32. 
о 1 ^ 2 ^ 

Abdomen greatly stretched at the groins and most of the thigh 
free. Upper lip highly cleft and serves as a prehensile organ. Stom- 
ach complex ("ruminant") and consists of four chambers, but differs 
somewhat compared to the stomach of true ruminants (the third 
chamber is not distinct while the first is designed to store water). 

'As this suborder is not found wild in the Soviet Union, its characteristics are very 
briefly summarized. It is possible that the modern species of wild camel {Camelus hoc- 
trianus, not as widespread as the Pleistocene Camelus knoblochi) survived in Central Asia 
and Kazakhstan until recent times. It is known from the kitchen-refuse of the Andronov 
civilization (1,000 to 1,500 B.C.; Vereshchagin, 1956). 



84 

Chewing the cud is characteristic. Cecum short. Skin glands few 
(occipital glands). Hair cover normal, well developed, sometimes 
even extremely well developed. Functional teats two and inguinal in 
70 position. Preputial orifice set far behind and turned backward. Pla- 
centa diffuse. In two species one or two large humps occur on the 
back. Red coфuscles oval (the only case among mammals). Sex- and 
age-related changes absent. 

These are desert, steppe, and mountain animals; they form herds 
and are strictly herbivorous. 

As wild animals they are distributed in Central Asia and South 
America; the domesticated form is found in the arid zones of Asia 
and North Africa (camels), and South America. 

Tylopoda are a very distinct group with several primitive charac- 
teristics and some specialized and progressive characteristics. They 
became isolated in the Middle Eocene and the present-day family of 
camels (Camelidae) has existed from the Upper Eocene onwards. Of 
the several subfamilies and genera, only one subfamily (Camelinae) 
with two genera, the true camel (Camelus L., 1758) and llama {Lama 
Fisch., 1775), has survived to date. 

The suborder has four living species: the two-humped camel 
{Camelus bactrianus L.) well known in the wild state in the deserts 
of Central Asia (almost exterminated at present), the one-humped 
camel {Camelus dromedarius L.), the guanaco {Lam.a guanicoe 
Mill.), and the vicuna (L. vicugna Mol.) inhabiting South America. 

The one-humped camel was never known in a wild state even in 
historic times. Only feral camels existed in North Africa. According 
to one view the one-humped camel is not a distinct species but only 
a domesticated, more civilized form of the two-humped animal. 
Both forms interbreed freely. In the embryonic stage the one- 
humped camel has two humps. At the same time fossils of one- 
humped camels are known from the Tertiary formations of North 
Africa and used to live in a domesticated state with inhabitants of 
the Sahara even in the Stone Age. 

The two-humped camel is a domesticated animal in Central Asia 
and Kazakhstan, while the one-humped animal is known in North 
Africa, Asia, Turkestan, Australia, and to some extent North Amer- 
ica. The llama {Lama glam.a L.) is a domestic form of guanaco, 
while the alpaca {Lam.a pacos L.) is a domestic form of vicuna. 

Feral one-humped camels are found in the estuary of Guadal- 
quiver [Spain] (?) and in the United States (Texas, Arizona, and New 
Mexico).* (V.H.) 

*No longer present in the United States — Sci. Ed. 



Suborder RUMINANTIA Scopoli, 1777 



71 Ruminantia are specialized artiodactyls of small, medium, or large 
size, with relatively long limbs, and mostly of light and slender 
build. 

These animals are hooved, with four or two digits and typical 
hooves. Lateral digits, if present, rudimentary and, in most cases, do 
not reach the ground and are nonfunctional. With few exceptions 
(Tragulidae), metatarsal and metacarpal bones of lateral digits 
either rudimentary and do not articulate with the tarsal and carpal 
regions, or completely absent. With one exception,' typical cannon 
bone present with two parallel, blocklike articular surfaces at the 
distal end. Carpal and tarsal bones fuse to some extent (at least the 
cuboid and naviculare are invariably fused). Ulna invarirbly 
reduced, set behind radius and in contact with it, or sometimes the 
two bones fuse; exceptionally (Tragulidae) ulna fully developed and 
separate. Shaft of fibula reduced (only the rod-shaped central por- 
tion sometimes preserved) and only the rudiments of its proximal 
and distal ends present (it forms an independent bone, os malleo- 
lare). As an exception, in Tragulidae, shaft of fibula complete. Trans- 
verse processes of cervical vertebrae pierced by vertebrarterial canals 
(art. vertebralis). 

Skull with greatly or moderately elongated facial part except in 
diminutive species; considerable development of cranial region 
("infantilism" in diminutive species). Longitudinal axis of skull 
usually highly flexed. Orbit closed from behind. Ethmoid pit well 
developed. Tympanic bulla tubular; exceptionally (Tragulidae) 
with lamellar osteosepta. Sagittal crest absent. Horns (several types) 
usually present at least in the male but often in both sexes; absent in 
a small number of species. Condyle of lower jaw elongated trans- 
versely (permitting lateral movements of lower jaw while rumi- 
nating). 

Upper incisors absent. Horny plate occurs on the lower surface 
of the anterior part of premaxillae. Upper canine absent in most 
species but present in some (in hornless forms — Tragulidae and 
some Cervidae) where it is highly developed in males. First upper 

^Hyemoschus (Tragulidae). Even in this species partial or total fusion of the 
metatarsals takes place in old age. The progressive specialization of limbs in artiodac- 
tyls generally occurs first and more intensely in the hind limbs which, evidently, is 
associated with their more dominant role in locomotion oDmpared to the forelimbs. 



86 

premolar normal (does not have the form of a canine) and not separ- 
72 ated by a diastema from row of cheek teeth. Lower incisors flat and 
broad (spatulate). Lower canine in contact with incisors and very 
similar in form. Cheek teeth cusps crescent-shaped (selenodont); 
curvatures of crescents in upper and lower jaws set opposite to each 
other. Molars and premolars hypsodont to various degrees, usually 
with late closing roots and sometimes passing into prismatic type 
(Tragulidae and Cervidae), and brachyodont. General dental 
formula: 

i|, с -^^, pm |, m| = 34 to 32. 

Stomach complex ("ruminant") with four well -developed sec- 
tions, three as an exception ("third stomach" rudimentary) in Tra- 
gulidae. Rumination of regurgitated food and marked participation 
of bacteria and protozoa in digestive process are characteristic fea- 
tures. Large cecum invariably present. In most species integumen- 
tary glands well developed on head (facial and occipital), legs 
(carpal, tarsal, and metatarsal), inguinal region, and some other 
parts. As an exception (Tragulidae), integumentary glands (at least 
facial, tarsal, and metatarsal) absent. Hair coat normal (bristles 
absent) but partly reduced in some cases. Teats inguinal and two or 
four in number. Sexual dimorphism usually well manifested, some- 
times very sharply, and occasionally indistinct. Age related changes 
sharp in some but relatively weakly manifested in most. Placenta 
cotyledonary; diffuse as an exception in Tragulidae. 

Ruminants are an extremely diverse group of artiodactyls. They 
include animals varying widely in size, external appearance, and 
biology such as mouse-deer, musk deer, deer, giraffe, ox, goat, sheep, 
and diverse varieties of antelope. The largest of these forms, the 
giraffe (see characteristics of the order), attains a weight of 1,200 kg 
and the smallest weighs 2 to 3 kg. 

With few exceptions (ox), ruminants are slender and mobile 
animals capable of sustained fast running (fastest of all nonflying 
mammals); some are adapted to climbing rocks. They inhabit 
extremely diverse biotopes, from tundras to tropical forests and 
marshes, most of which are associated with open biotopes (steppes 
and steppelike zones and mountains); a few are confined to forests 
and marshes or dense grassy thickets (similar to hares). These 
animals live in herds to some extent, except Tragulidae which live 
singly. They give birth to one or two offspring; four to seven are 
exceptional (Hydropotes). They are herbivorous and feed mainly on 
grassy vegetation. 



87 

Ruminants are a very widely distributed group of the order, 
inhabiting all of its geographic range except Tierra del Fuego, 
Madagascar, and the West Indies. The southeastern boundary of 
their range extends along the island region between Asia and Aus- 
tralia and includes the islands of Timor, Ceram, Buru, Halmahera 
(Dzhilolo), and the entire group of Moluccas and the Philippines. 
Ruminants are absent in Tanimbar, Kai, Aru, and New Guinea if 
sambar, acclimatized there, are not taken into consideration. Several 
species (deer, tahr, chamois, and others) have been acclimatized in 
New Zealand. 

In diversity of adaptation, biological types, distribution, number 
of species, and numerical strength Ruminantia are an extremely 
flourishing extant group of artiodactyls and of hoofed animals in 
general. Although in some morphological features they are no more 
specialized (often less) than Tylopoda, on the whole (biologically) 
they represent a most progressive branch of the order. 

Present-day ruminants, in spite of their varied diversity, are 
extremely distinctive and sharply delineated from suborder Suiformes. 

The differences between suborders Tylopoda and Ruminantia 
are less striking. This has prompted some zoologists to combine 
these two groups into a common suborder, which is hardly justifia- 
ble. Both groups possess important differences; they flourished inde- 
74 pendently from the Eocene period (appearing at the commencement 
of evolution of the order). The most striking similarities between 
Tylopoda and Ruminantia (complex stomach, some structural fea- 
tures of limbs, etc.) are largely the consequence of their parallel 
adaptation. There are no "connecting links" whatsoever between 
the two groups. Nevertheless, Ruminantia are closer to Tylopoda 
than to Suiformes. 

In basic structural plan the suborder represents an extremely 
homogeneous group. As can be seen from the morphological char- 
acteristics given above, only the mouse-deer (Tragulidae) with its 
several primitive features stands out prominently in the suborder.^ 

The ruminant group Tragulina appears in the Upper Eocene 

^Some specialists in taxonomy are inclined to separate them from the remaining 
ruminants, i.e., not only typical ruminants (Pecora, i.e., Ruminantia without Tra- 
gulina) but also Tylopoda, as an equivalent group (usually "sections") within the 
same suborder. This combination and some others which assign to the mouse-deer an 
even greater independent status is irrational. In spite of all their distinct characteris- 
tics, Tragulidae cannot be regarded as equivalent to Tylopoda and even less so to 
Suiformes. Additional characteristic features of Tragulidae are a very thin interocular 
septum and the fusion of the optic foramena into a single opening. 



89 

(Europe, North America, and Asia). This group had evolved mostly 
in the Oligocene at the close of which its extinction commenced; 
however, an extremely small number of species have survived in the 
Recent epoch as relicts. Furthermore, Tragulina were evidently 
never particularly abundant. Pecora occur only in the Lower Oligo- 
cene (deer; Asia) and some groups even in the Lower or Middle 
Miocene (giraffe and Bovidae). Tragulina during the period of their 
greatest development evidently served as the parent group for 
Pecora. After the Miocene there was an exceptionally rapid develop- 
ment of Pecora, which became an extremely divergent, abundant, 
and flourishing group of artiodactyls (and ungulates in general), 
that survived until the end of the Tertiary period and flourish even 
now. This is particularly true of Bovidae. The remaining groups of 
Pecora developed relatively less evenly or recorded no special devel- 
opment whatsover; they have been reduced at present to groups 
wfiich are becoming extinct (Antilocapridae and Giraffidae). 

Until recently the taxonomy of supraspecific categories of rumi- 
nants (Pecora) was extremely imperfect". At present, however, the 
basic features of the supraspecific categories can be regarded as fairly 
well understood and generally accepted. Some generic classifications 
of deer, more so the division of Bovidae into subfamilies (wfiicfi 
until recently remained extremely artificial and arbitrary) constitute 
exceptions. 

The taxonomy of various species of the suborder is relatively 
simple, but until recently had been studied little and not always 
along correct lines. The actual picture of the species composition of 
this group (especially of African forms) was understood only in the 
last decade. This led to a sharp (and as yet incomplete) reduction in 
number of species. 

The suborder has been divided into two infraorders — Tragulina 
and Pecora. In toto the suborder comprises 255 genera (about 61% of 
all artiodactyls) placed under 15 families (60% of artiodactyls). Of 
these, 76 are extant (about 89% of extant artiodactyls), representing 5 
families (about 55% of extant artiodactyls), and embracing about 132 
species (about 88% of all extant artiodactyls). 

Tragulina are classified into three superfamilies of which two 
are extinct and one extant — Traguloidea — comprising one extinct 
and one extant family — Tragulidae. The latter includes two genera 
(Tragulus and Hyemoschus; about 3% of genera in the suborder) 

'Partly due to vague ideas about species differentiation, Pecora until recently have 
been subjected to extremely diverse, often peculiar, taxonomic divisions. Almost every 
genus of deer has been raised to the rank of family by one taxonomist or the other. 



90 

with four species (three in southeast Asia and one in western Equa- 
torial Africa; about 2.7% of species of the suborder). 
75 Infraorder Pecora is divided into three superfamilies: Cervoidea 
(solid-homed or deer) with a single family (Cervidae), Giraffoidea 
(giraffe) with one extinct and one extant family (Giraffidae; Africa), 
and Bovoidea (hollow-horned) with two families — Antilocapridae 
(pronghom; North America) and Bovidae (hollow- horned or oxen). 
Family Giraffidae includes one extinct and two extant subfamilies: 
Giraffinae (one genus, Giraffa, with two species) and Palaeotraginae 
(one monotypic genus, Okapia). Family Antilocapridae includes 
one extinct and one extant subfamily — Antilocaprinae with one 
monotypic genus (Antilocapra). 

Pecora comprise 74 Recent genera (over 97% of genera of the 
suborder) and about 128 species (97% of species of the suborder). 

Many species are African in origin and fewer associated with 
Asia. In North and South America ruminants are few and poorly 
represented. 

As the most abundant species of the suborder, ruminants are of 
utmost economic importance in all the features listed above (see 
characteristics of the order). 

Fauna of the Soviet Union include one (Pecora) of the two 
infraorders, two (Cervoidea and Bovoidea) of the four superfamilies, 
and two (Cervidae or deer and Bovidae or hollow-horned animals) of 
the five families. The 18 species account for about 13% of species of 
the suborder or about 6% of mammalian species of the USSR. On the 
whole the group is not abundantly represented in the Soviet Union 
but is quite diverse. 

Ruminantia are valuable game animals in the Soviet Union, 
providing important products for local consumption. (V.H.) 



''Data on the composition of groups represented in Russian fauna are presented 
later under appropriate sections. 



Infraorder PECORA Linnaeus, 1758 

Superfamily CERVOIDEA Simpson, 1931 
Family CERVIDAE Gray, 1821 

76 Ruminants of moderate, smalls or rarely relatively large size, with 
long legs; most are slender and light in build. 

With some exceptions digits four. The lateral ones carry small 
hooves and usually have skeletal elements (phalanges); exception- 
ally they are relatively well developed and under certain conditions 
are of some importance in the movement of the animal. The meta- 
tarsal bones of digits II and V are completely reduced and only the 
upper or lower parts of the corresponding metacarpal bones are 
retained. 

Facial part of lacrimals very well developed and, with few excep- 
tions {Moschus, Capreolus, and Pudu), contain pits (depressions) 
for preorbital scent glands. Orbital region of lacrimals penetrated by 
two (exceptionally one) tear ducts. Ethmoid pit well developed and 
sometimes very large. Jugal greatly extended in facial part of skull; 
lateral surface of maxilla correspondingly shortened. Due to brachy- 
odont dentition alveoli on maxilla shallow. Pneumatization of skull 
bones poor (lacrimal, maxilla, and anterior part of frontal). In forms 
with antlers, because of their growth, frontals usually greatly 
extended backward and parietals very much reduced. Orbit large and 
turned sideways. Premaxilla relatively insignificant in size. Tym- 
panic bulla loosely articulate with skull, usually small, always tubu- 
lar in shape, and generally with an uneven external surface. 

In most species (exceptions, Moschus and Hydropotes) males, 
and in one case (Rangifer) females also, have antlers which are 
nonpneumatic projections (apophyses) of the frontals (not on separ- 
ate bony formations called os cornu). Antlers consist of bony (not 
horny) material (mesodermal) and are formed as processes (conti- 

77 nuations) of apophyses of the frontals; they are shed once a year and 
grow again. During the period of growth antlers are soft and 

Some researchers hold that the skin also participates in the formation of antlers 
and thus regard them, to some extent, as compound structures, not exclusively skele- 
tal. In any case there can be no dispute about the basic differences in the formation 
and structure of antlers of deer and hollow-horns. 

^The biannual shedding of antlers seen in Elaphurus davidianus is evidently the 
result of prolonged domestication. ^ 



92 

covered with a hairy skin ("velvet"), which during antler ossifica- 
tion dies and sloughs. Antlers, with few exceptions (Elaphodus, 
Mazama, and Pudu), are branched, i.e., consist of a main beam with 
tines; fully developed among adults of various species, normally 2 
(Muntjacus) to 20 tines occur." At its base an antler exhibits a promi- 
nent ring ("rosette") with an uneven surface (channels through 
which blood vessels feed the antler during the period of growth). 
Cheek teeth, at least the anterior ones, brachyodont. Roots, com- 
pared to size of crowns, small. Lateral vertical ridges on molars 
joined by horizontal ridge at base of crown, on boundary of its 
alveolar edge. Upper canine usually reduced; may, however, be 
absent altogether, or well developed, or even highly developed (par- 
ticularly in males). Growth of canines related to development of 
antlers (particularly large in antlerless forms), quite prominent in 
species with poorly developed antlers, and rudimentary in forms 
with highly developed antlers. Dental formula: 

i , с ^ , pm I , m I = 32 to 34. 

Integumentary glands well developed and diverse. Hair short 
and densely set. Color of adults uniform, dark, or sometimes with 
light-colored spots. In many species light-colored area ("speculum") 
present on rump. Sexual dimorphism prominent in majority of 
species; age-related changes also distinct. Spotted coloring of young 
animals (in large number of species) characteristic feature. Gall 
bladder, with few exceptions, absent. Placenta with few cotyledons. 

Deer constitute a homogeneous family in which only a few spe- 
cies deviate markedly from the general pattern. These differences are 
not significant per se but associated with the absence or underdevel- 
opment of antlers and a small body size (relatively large and com- 
plex antlers present in larger species). On the whole, the same set of 
characteristics are consistently seen throughout the family. The 
largest of the species (moose from Kenai, Alces alces gigas) attains a 
body length of 310 cm, height at withers 233 cm, and weight 770 to 
815 kg (Anthony, 1928).^ The smallest of the species have a body 
length of about 90 cm, height at withers about 51 cm, and weight 9 
to 10 kg (Hydropotes) or even less (Muntjacus reevesi has a height of 
40 to 45 cm at withers and Pudu 35 cm, i.e., the size of a large fox). 

Antlers of some races of C. elaphus witfi 32 or more tines represent a rare and 
extreme individual variation. 

"These data, if not exaggerated, must refer to a very large specimen. 



93 

This group consists of mainly forest and partly forest-steppe 
species and a few tundra species. On the mountains they are found 
up to tfie upper limit of forests, entering into tfie lower regions of 
the alpine zone. Brandies of trees and various species of shrubs form 
the major part of tfie diet of many species. 

Deer have a relatively wide distribution. They inhabit tfie wfiole 
of South and Central America except for Tierra del Fuego and the 
West Indies, tfie whole of North America including ttie Arctic 
archipelago, tfie wtiole coastal belt of Greenland except tfie east 
coast, and Unimak Island (Aleutian Islands). In tfie Old World tfie 
geographic range covers northwestern Africa," the wfiole European 
and Asian mainland up to and including Palestine, tfie Syrian part 
of United Arab Republic, and Mesopotamia in tfie south. In ttie 
nortfi, the range covers Spitsbergen, Novaya Zemlya, southern 
79 islands of Severnaya Zemlya, and tfie New Siberian Islands. In the 
south the range extends up to Ceylon and all of tfie island region 
between Asia and Australia in tfie east and southeast including 
Timor, Seram, Halmahera, and tfie Pfiilippines. The range also 
covers tfie east Asian islands, except tfie Kuril and Bering Sea 
Islands. 

Extant forms of deer represent a very distinct, well-defined group 
of ruminants. In spite of sfiarp differences between extreme forms, 
there are no transitional forms between deer and otfier families. The 
peculiar structure of fiorns and tfieir sfiedding in Antilocapra 
(prongfiorn) are essentially unusual features of purely superficial 
similarity. 

The family is, to a high degree, homogeneous. Numerous attempts to 
subdivide it into several families (see above) lack sufficient basis and 
have now been abandoned. Only musk deer {Moschus), which even 
now are sometimes regarded as a distinct family, stand apart promi- 
nently within the family. 

'At present the geographic range of deer (C. elaphus) has shrunk very much. 
Formerly it extended into Algeria, Tunisia, and Morocco fairly extensively, but deer 
are now foundonly in a small section at the edge of Algeria and Tunisia (Heim de 
Balzac, 1936). Fallow deer (C dama) were acclimatized in Egypt even in the distant 
past and are not a primitive species there. However, some data point to fallow deer 
being autochthonous in this region. 

Deer (Rusa complex) were acclimatized in New Guinea and New Caledonia. 
Even before the appearance of Europeans, the local inhabitants evidently raised deer 
on some islands. It is possible that юте islands within the area under description 
were colonized thereby and the boundary described therefore may not be altogether 
natural. It is, however, difficult to establish it now. Probably, artificial colonization 
also influenced the development of some morphological characteristics throughout 
the island zone. This is true to an even greater extent of swine. 



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95 

Cervidae appeared in the Lower Oligocene (Asia) as relatively 
poorly differentiated forms, comprising the subfamily Palaeomery- 
cinae, which became extinct in the Pliocene. This family is clearly 
linked with Oligocene Traguloidea and originated from them. Deer 
evolved from very small antlerless forms. One of the features of their 
evolution has been the acquisition and increase in size of antlers, 
which have become increasingly complex in the course of time. 
During their evolution no notable diversity developed in the deer 
family and they were not particularly abundant. They evolved rela- 
tively uniformly and at present, although not very abundant (fewer 
than Bovoidea), they represent a flourishing and extensively distri- 
buted group. The geographic range and population of deer, like 
those of all other large mammals, are at present diminished due only 
to man's intervention. 

The family structure is simple. The time for further division into 
families and subfamilies and their separation into innumerable gen- 
era is well past, although the division of deer into various genera 
cannot yet be regarded as complete. Marked progress has been made 
in the separation of various species. The old method of intense 
subdivision and narrow interpretation of species is no longer 
acceptable, but partly still implemented for southeast Asia, particu- 
larly in the island region. This is largely explained by the develop- 
ment of complex structures in the animals of these regions. 

The family is divided into seven subfamilies, two extinct 
(Palaeomerycinae and Dromomerycinae) and five extant: Moschinae 
(musk deer), Muntjacinae (barking deer), Hydropotinae ("water 
deer"), Cervinae (red deer or plesiometacarpal deer), andOdocoilei- 
nae (moose, Virginia deer, and related groups — telo metacarpal 
deer). The family comprises 50 genera (about 19% of all ruminants). 



'The classification presented here corresponds in its main features to that pro- 
posed by Simpson (1945), which is now widely adopted. One departure, recognition 
of the separate subfamily Hydropotinae, is supported by several researchers (Pocock, 
1923; Allen, 1940; Ellerman and Morrison-Scott, 1951). 

None of the prevailing schemes are very satisfactory since the subfamilies adopted 
are not of the same value in taxonomic importance. Strictly speaking, only two 
groups are distinctly identifiable within the family. One is Moschinae and the other 
includes all the remaining members of the family. On the other hand, there is no 
doubt that if subfamilies Muntjacinae, Cervinae s. str., and Odocoileinae are taken 
into consideration, linking Hydropotes to Odocoileinae is not in order, but the 
former should be isolated in the same manner as the other groups indicated above. 
Evidently, it would be more natural ultimately to subdivide the family into two 
subfamilies (Moschinae and Cervinae) and place the rest into a single suprageneric 
group, but of a much lower rank (tribe). ^ 



96 

of which 17 are extant (Simpson, 1945)^ (22 to 23% of present-day 
ruminants). The total number of species is 36,^ i.e., about 24.3% of 
species of the suborder. 

Cervidae are game animals providing meat, hide, often fur 
(mostly used by local inhabitants), and pharmaceutical material 
(European and Chinese medicine — antlers in velvet, tails, sex 
organs, and musk glands). Antlers are used for utilitarian and decor- 
ative articles; they are also valued as trophies. In the last century the 
population and range of most of these species were greatly reduced. 
One species (Elaphurus davidianus) has become extinct in its native 
land (China) with only several dozen animals preserved in zoos 
(England and Australia). Others face the threat of extinction and 
several survive under protection. 

At some places deer are objects of intense hunting. Several spe- 
cies have been acclimatized outside their natural geographic range 
as game and ornamental animals (fallow deer in Europe and South 
America, sambar in Mauritius and Rodriquez, a few forms in New 
Zealand, reindeer in South America, and so on). One species (rein- 
deer) exists in both wild and domesticated states. The degree of 
domestication is not very high. Experiments are underway for the 
domestication of moose. Some deer are held in a semidomesticated 
state in special farms for procuring their antlers in velvet (sika, 
maral, and Manchurian wapiti) or raised in zoos (fallow and Pere 
David deer). 

The Russian fauna includes three of the five'° subfamilies 
(Moschinae, Cervinae, and Odocoileinae), five" of the 13 genera 
(Moschus, Ceruus, Capreolus, Alces, and Rangijer) {гЬот 36% of 
extant genera), and six of the 36 species (about 16% of extant species). 
Deer constitute about 1.5% of species of Russian fauna. 

In the Soviet Union members of the family are found throughout 
the territory except in steppe and desert zones. In the later zones they 
occur only in a few intrazonal points which are favorable for 
habitation. 



^Eleven (Flerov, 1952) or 13 (Haltenorth and Trenze, 1957) is much closer to the 
actual number. Within the family the categories of subgenera are well substantiated. 
The list of genera given below should be referred to when discussing subfamilies. 

'According to the data of Haltenorth and Trenze (1957). Lydekker (1915) recog- 
nized 48 and Flerov (1952) 50 species. These numbers are clearly exaggerated. 

'"Muntjacinae comprise two genera {Muntjacus and Elaphodus); a few species 
occur in south and southeast Asia. Hydropotinae (one monotypic genus) is found in 
eastern China and the Korean Peninsula. 

^^Dama is considered a subgenus. Moreover, fallow deer is not an indigenous 
species of the Soviet Union; it was acclimatized as a game animal. 



97 

The economic importance of deer in the Soviet Union has 
already been discussed. However, no intense raising of these species 
for use as game animals has been undertaken. Deer are also con- 
served in several preserves. Hunting of some species is totally or 
partially prohibited. (V.H.) 

Key to Species of Deer (Cervidae) 

1 ( 2). Antlers absent in males and females. Upper canines very 
large: in adult males project out of mouth; in females their 
length is greater than height of molars. Preorbital glands 
absent. Hind limbs almost 30% longer than forelimbs. 
Lacrimal vs^ithout depression. Usually only one tear duct 
present. Body length of adult animals not more than 110 cm 

and maximum skull length 165 mm 

Musk deer, Moschus moschiferus L. (pg. 101) 

2(1). Antlers present in males and sometimes in females. Upper 
canines absent or poorly developed; their length not more 
than height of molars. Preorbital glands present and often 
well developed. Hind limbs only slightly longer than fore- 
limbs. Lacrimal with depression, often large and deep. Tear 
ducts invariably two.. Body length of adults exceeds 110 cm 
and maximum skull length more than 180 mm. 
81 3 ( 6). Muzzle and area between nostrils entirely covered with hair or 
with only a very small bare patch on upper lip between nos- 
trils (most of area between nostrils covered with hair). 

4 ( 5). Space between nostrils entirely covered with hair; muzzle not 

enlarged. Vomer divides nasal passage so that pair of passages 

formed. Females with antlers 

Reindeer, Rangifer tarandus L. (pg. 430) 

5 ( 4). Between nostrils small section of bare skin occurs but much 

of it covered with hair; muzzle of adults very much enlarged. 
Vomer does not divide nasal passage and hence latter not 

paired. Females without antlers 

Moose, Alces alces L. (pg. 307) 

6(3). Space between nostrils, sometimes neighborhood of nostrils, 
and entire muzzle bare. 

7 ( 8). Tail very short, does not project out of fur, and not discerni- 

ble externally. Preorbital gland poorly developed and pit half 

diameter of eye. Depression on lacrimal small and faint 

Roe deer, Capreolus capreolus L. (pg. 243) 

8 ( 7). Tail well developed, projects out of fur, and invariably very 

distinct. Preorbital gland large (equal to or slightly less than 
diameter of eye). Depression on lacrimal large and deep. 



98 

9 (10). Tail relatively short; length (together with terminal hair) less 
than ear length. Tail not black at base. Speculum large and 
ocherous or with an ocherous tinge, but not pure white. Body 
size large; height of adult male at shoulders usually not less 
than 120 cm and maximum skull length exceeds 350 mm. 
Summer coat without spots or with stray spots along sides of 
backbone and top of back, usually in females. Antlers not 
flattened and with two tines in lower part of beam (brow and 
bez) Red deer, Cervus elaphus L. (pg. 159) 

10 ( 9). Tail relatively long; length (together with terminal hair) 

more than ear length or almost equal to it. Tail with black, 
white fringed zone at base. Speculum white. Summer coat 
with bright, light-colored spots on back and sides. Only one 
tine (brow) at base of antlers. 

11 (12). Body size, small; height of adults at shoulders up to 1 m or so. 

Tail markedly longer than ear. Hair around urinogenital ori- 
fice long and penicillate in arrangement. Upper canines 
absent. Antlers flattened at ends and shaped like a vertical 

shovel. Maximum skull length under 300 mm 

Fallow deer, Cervus dama L. (pg. 981) 

13 (12). Body size slightly larger; height at shoulders over 1 m. Tail 
nearly equal to ear length or slightly longer. Hair around 
urinogenital orifice not long and not penicillate in arrange- 
ment. Anders not flattened at ends and not shaped like a 

shovel. Maximum skull length over 300 mm 

Sika deer, Cervus nippon Temm. (V.H) (pg. 131) 

Subfamily MOSCHINAE Zittel, 1893 (Musk Deer) 

Size, small. 

Toes four. Lateral ones relatively well developed with normal 
phalange skeleton; only distal portions of metacarpal and metatar- 
sal bones of toes II and V present, while proximal portions reduced 
("telometacarpal" ends). Lacrimal without depression for preorbital 
82 gland. Orbital region of lacrimal pierced by single tear duct. Nasal 
process (pr. nasalis) of premaxilla reaches, or almost reaches, fron- 
tal, or only separated from it by ethmoid pit. Ethmoid pit small. 
Parietals normally developed (not reduced). Antlers and apophyses 
of frontals absent. 

In males upper canine extremely well developed and projects out 
of closed mouth; it is sharp, points directly downward, and grows 
throughout life. Females invariably with canines but they are small. 
Dental formula: 



99 



3 ' 1 



3 
pm н- 



m I = 34. 



Hind limbs markedly longer and more massive than forelimbs, 
giving animal an altogether characteristic appearance — posterior 
part of body notably more massive than anterior, very much raised, 
and locomotion saltatorial. Exposed anterior portion of muzzle 
large and surrounds nostrils. Tail very short. Preorbital and metapo- 
dial glands absent. Caudal glands and large musk gland on abdo- 
men (in preputial region) present. Glands also present on outer 
surface of thighs (Yu.A. Salmin); glans penis with filiform protuber- 
ance. Gall bladder present. Teats two. 

Molt occurs once a year. 

Moschinae are montane forest (probably often alpine) forms 
associated with rocks and steep inclines. 

Found in the Himalayas, China, including the northeastern 
part, the Far East, and eastern and central Siberia (see description of 
musk deer). 

Musk deer, as mentioned in the family description, represents a 
distinctly different group of extant deer and their status as an inde- 
pendent subfamily is indisputable. Until recently some researchers 
have long been inclined, without much basis, to separate them com- 




Fig. 26. Geographic range (reconstructed) of subfamily and genus of musk 

deer, Moschinae and Moschus L., and range of musk deer species Moschus 

moschiferus L. V.G. Heptner. 



100 

pletely from deer into a special family. Some even considered musk 
deer a connecting link between families Cervidae and Bovidae. This 
view, based on a few insignificant characteristics, has no adequate 
basis whatsoever, any more than attempts by some zoologists to place 
this group among Traguloidea. 
83 Primitiveness of Moschinae among extant Cervidae is indubita- 
ble. Fossils of musk deer are known only from the Pliocene and 
probably Lower Pliocene of Asia, however, and even these belong to 
the Recent genus. 

The subfamily is represented by only one genus (about 8 to 9% of 
extant genera of the family) with one species (about 3% of species of 
the family).'" 

Musk deer are caught for musk, which is used in the perfumery 
industry. 

There is only one species in the subfamily, constituting about 
0.3% of species of Russian fauna. 

In the Soviet Union musk deer are distributed in central and 
eastern Siberia and in the southern part of the Far East. (V.H.) 

Genus of Musk Deer 

Genus Moschus Linnaeus, 1758 

\lb%.Moschus. Linnaeus. Syst. Naturae. Ed. X, vol. I, p. 66. 

Moschus moschiferus Linnaeus (Opinion 75 of the International 

Committee on Nomenclature). 
1848. dontodorcas. Gistel. Naturgesch. d. Tierreichs, p. 82. 

Moschus moschiferus Linnaeus. See characteristics of the 

subfamily. 



'^Suggestions that the genus contains three species — Himalayan M. chrysogaster 
Hodgson, Szechwan M. berezovskii Flerov, and Siberian M. moschiferus Linn., orM. 
moschiferus, M. berezovskii Flerov, and M. sibiricus Pall. — according to another 
nomenclature (Flerov, 1930 and 1952)— have not been favorably received. Neverthe- 
less, the extremely small Szechwan musk deer should be studied further from mor- 
phological and geographic points of view. To date only two typical individuals have 
been caught from the region where the much larger M. m. sifanicus Biich. lives. 
Prevailing opinion maintains that these two individuals are no more than extreme 
variants or even simply abnormally developed individuals ("starvelings") (Allen, 
1940). 



101 
MUSK DEER 
Moschus moschiferus Linnaeus, 1758 

\lb%.Moschus moschiferus. Linnaeus Syst. Naturae, 10th ed., vol. 1, p. 

66. Altai." 
1779. Moschus sibiricus. Pallas. Spicil. zooL, vol. 13, p. 29. Stanovoy 

mountains. 
ISSO. Moschus altaicus. Escholz. Isis (Oken), p. 106. Mongolia. 
\9ll. Moschus parvipes. Hollister. Proc. Biol. Soc. Washington, vol. 

24. Mountains at Мокро in southern Cholla province, South 

Korea. 
\92^. Moschus moschiferus arcticus. Flerov. Doklady Akad. Nauk 

SSSR, p. 516. Mountains of Tulayakh-khaya, 460 km north of 

Verkhoyansk (northeastern branch of Tas-Khayakhtakh). 
1928. Moschus moschiferus sachalinensis. Flerov. Ibid., p. 517. 

Aleksandrov, Sakhalin. 
1954. Moschus Moschiferus turovi. Zalkin (Tsalkin). Doklady Akad. 

Nauk SSSR, vol. 46, no. 8, p. 366. Terney Bay in Sikhote-Alin 

sanctuary, Ussuri region. (V.H.) 

Diagnosis 

Moschus moschiferus is the only species of the genus. 

Description 

Musk deer in general appearance differ sharply from other deer and 
all other ungulates. 

In build musk deer are slender and elegant. Their hind limbs 
are very long and well muscled; their forelimbs are relatively 
short, thin, and weak; and their chest usually small. Hence the hind 
quarters appear considerably larger, wider, and more massive than 
the anterior part of the body, the back highly arched, and the animal 
much higher at the sacrum than at the shoulders. With its small 

'^Linnaeus distinguished "Tataria versus Chinam" forms. The nearest part of 
"Tataria" in the direction of "China" where musk deer are raised is Ahai. This was 
accepted by several researchers and authors of monographs, including Soviet authors 
(Flerov, 1930; Tsalkin, 1947). There is no justification for altering this established 
view to regard "northern India" (Himalayas) as the type locality (Flerov, 1952). 
Furthermore, to do so would lead to needless difficulties in nomenclature (see Flerov, 
1952). 

'"Stanovoy mountains are considered the type locality by Ellerman and 
Morrison-Scott (1951), probably based on the writings of Chaworth-Musters. 
Initially, Flerov designated "eastern Siberia" the type locality (1928 and 1930) but 
later opted for Altai (1952). 



102 

head and relatively thin, short, and fairly low neck, a musk deer 
looks "mournful" or "frightened". Even at a slow pace the animal 
moves with fairly long, extremely easy, soft, springy jumps, with its 
head lowered, emphasizing all the more its characteristic appear- 
ance. The above proportion of limbs and some other characteristics 
(elongation of head of femur, straightened pelvis, and others) 
designed for saltatorial movement are developed much more in 
musk deer than in ^ny other ungulate, including even typical mon- 
tane animals, and are undoubtedly advantageous for living in 
mountains and hills. 

Head relatively small and elongated; eyes fairly large; ears long 
and broad, with fairly rounded tips, and extremely mobile. At tip of 
muzzle large bare patch of black coloration surrounds nostrils. 
Large, somewhat saberlike, fine, and extremely sharp canines pro- 
ject directly downward from mouth of male. Tips of canines in old 
males descend considerably below chin. Canines of females not vis- 
ible externally. Tail very small and does not project out of fur. In 
adult males tail bare (only tuft of hair present at tip), flat, thick, with 
rounded end. In females and young males tail thin and covered with 
hair. Hooves of middle toes sharp, narrow, and drawn out. Lateral 
hooves relatively large, only slightly smaller than medial ones, 
sharp, and elongated; ends reach base of hooves of medial toes. 
Functionally, lateral hooves more active than similar structures in 
other Russian species of the family; in a standing animal they 
almost invariably reach the ground and are usually seen in 
hoofprints. 

Fur consists of hair of several types. Much of body covered with 
long, somewhat wavy, coarse (up to 0.5 mm in diameter), and elastic 
hair which is highly pneumatic and hence quite brittle. Length of 
hair on sacrum reaches 95 mm and at withers 65 mm; on sides 
sometimes shorter, and even more so on neck and abdomen, espe- 
cially on head and legs. Underfur relatively poorly developed, very 
soft, and delicate. Basal two-thirds of hair whitish and rest 
brownish-gray; tips dark brown or even blackish-brown. Some hairs 
with blurred, fairly broad, rust-colored band. 

The overall coloration of the body of adult animals in full winter 
coat is dark chocolate or even dark brown, and fairly uniform 
throughout the body. Head gray with an admixture of brown on 
top. Ears grayish-brown on back side, dirty gray inside, and tips 
blackish. Upper side of neck dark chocolate with grayish tinge. 
Against this background usually a yellowish band occurs contain- 
ing sometimes paired hazy spots. On lower side of neck a white or 
whitish band with an elongated medial brown patch extends from 



103 

lower jaw. Light-colored band runs farther onto chest and inside 
surface of forelimbs. Color of sides somewhat lighter than that of 
back. Light brownish-ocherous or rusty-yellow spots scattered on 
sides and along back, which usually form longitudinal symmetrical 
rows. Two rows occur on the front and sides and five or six on the 
thighs. Two rows usually extend along back, sometimes almost 
merging into longitudinal bands. Abdomen and limbs brownish- 
chocolate and front and inner side of legs grayish. 

Overall coloration of body exhibits fairly pronounced individual 
variations from light, almost yellow to brownish- black. Spots in 
85 particular highly variable. Usually large, bright, and distinct, but 
sometimes small, dull, and blurred. Sometimes obsolete on sides but 
invariably preserved on thighs. Contrarily, sometimes number of 
spots so great as to give an overall ocherous-gray tinge to much of 
trunk. There are no sex-related color differences, or such differences 
are slight; spotting in females appears to be somewhat more intense. 
Seasonal differences manifested in fur color; it becomes lighter 
toward the end of winter as a result of wear and fading. 

Molt occurs once and there is no summer coat as such. Com- 
mencing from the end of winter and nearly throughout the whole of 
the warm period of the year, old hair is lost and new commences to 
grow; this growth process ceases late in autumn. In summer the 
animal has short hair without underfur. Faded and worn remnants 
of old fur along with growing and already grown new hair are often 
seen on the back throughout summer (especially in nursing 
females). 

Age-related changes in color and hair coat are prominent. In 
new-borns hair is short and relatively soft. Background color dark 
brown. Entire upper side of trunk (back, scapular region, upper part 
of the sides, forelimbs, and thighs) densely covered with spots, form- 
ing a distinct light-colored "saddle cloth". Spots yellowish-white or 
rust in color and often placed so that longitudinal rows (sometimes 
transverse also) form, which often merge into bands. On the croup 
and thighs, where the band is larger, as many as five rows may occur. 
Large whitish patch occurs on head and under ears; inner surface of 
ears light ocherous and outer part rusty. Upper side of neck with two 
large, light rust-colored spots. Throat white with yellowish hue. 
Two light-colored bands run along lower side of neck onto chest 
and on inner side of forelegs, where they reach lateral hooves. In 
addition to whitish field on throat, two small white bands on each 
side extend backward along posterior side of cheek and throat. 
Abdomen white with a yellowish tinge; groins and inner surface of 
hind legs white. By winter the young animal molts into its first 



104 

winter coat, made up of coarse hair typical of adult animals, and 
resembling in color that of adults but with spots better developed. 
Second winter coat (at age 1.5 years) does not differ in color from 
that of adults.'^ 

Caudal glands located on upper surface of tail contain a secre- 
tion which exudes a goaty smell. Musk glands enclosed in a special 
sac of skin located in front of preputial orifice. In adult males sac 
may reach about 3 cm in width and 4 to 5 cm in thickness. Both 
categories of glands are associated evidently with sexual activity 
(probably for marking territory); structure and function of other 
glands (on the thighs) not known. Mammary glands bare and two 
teats 14 to 15 mm long located close together. 

Skull small with relatively well- developed cranial part and short 
facial part. Length of latter comprises 47 to 52% of maximum length 
of skull. Lacrimal flat and without depressions. Premaxillae with 
long nasal processes wedged between nasal and maxilla. Supraorbi- 
tal processes insignificant. Parietal crests converge behind and form 
an indistinct arrow-shaped crest. 

Teeth relatively small. Incisors and lower canines differ little in 
size and only slightly decrease in size from the center to the outside. 
Crowns flat and broad like leaves, fairly symmetrical (not bent to 
one side), and end ones somewhat superimposed on medial ones 
(canine on third incisor, third incisor on second, and second on 
first). Molars on lower jaw with well -developed additional tubercles 
on outer side. Upper canines of males highly developed and saber- 
like with very sharp tips. They are greatly compressed from the 
sides, the rear edge sharp and used for cutting, and the anterior 
surface more blunt. Length up to 70 or even 100 mm. Pulp cavity 
86 exposed and grows throughout life of animal or for most of it. 
Tooth fits loosely in alveolus and when in use can move slightly 
forward, backward, or sideways. This evidently prevents breakage to 
some extent. Upper canines present in females but small, although 
not smaller than lower canines, and do not project out of mouth. 

Sexual dimorphism in skull well manifested and associated 
primarily with varying degrees of development of canines. Entire 
frontal region of skull of males much more massive and broader 
than that of females. Sides of nasal region of male skull almost 
parallel, while in female converge forward like a wedge. Due to large 
size of alveolus of canine, premaxilla and maxilla in male large but 
nasal process of former lies almost horizontally. In females they are 

^^Views regarding much longer series of age-related changes and maturity of 
animals only in the third year (Flerov, 1952) are erroneous (as explained later). 



105 

almost vertical. Palate in region of canines broader in males than in 
females. Supraorbital processes and zygomatic arch in females 
weaker than in males, and overall skull size of females somewhat 
smaller. 

Age-related changes are more prominent in the skull of males 
than in females, due to progressive enlargement of canines with age. 
Consequently facial part becomes increasingly massive and enlarged 
in males, while in females it remains fairly uniform. 

Vertebral column intensely arched. Posterior part large and rela- 
tively massive, while anterior part much slighter in build. Lumbar 
region massive and equal to thoracic in length. Chest short and 
narrow (Flerov, 1952). Length of hind limbs more than that of fore- 
limbs (up to the shoulder joint). ^ 

Size and weight of animals undergo relatively few changes. This 
is primarily true of age-related changes, since growth and maturity 
proceed rapidly and the animal is already fully developed by the age 
of two years. There are no perceptible sex-related differences in size 
and weight. Newborns weigh 300 to 500 g, adult females in Sikhote- 
Alin up to 15 to 17 kg, and adult males slightly less (Yu.A. Salmin). 
Overall body length 86 to 100 cm, height at winters 56 to 67 cm, 
rarely 80 cm, height at sacrum 67 to 80 cm, length of ears 8 to 11 cm, 
and tail length 4 to 6 cm. (V.H.) 

Taxonomy 

Moschus moschiferus is the only species of the genus. 

Geographic Distribution 

Inhabitants of eastern Asia, almost from the northern forest bound- 
aries to southern China, Assam, and Burma. Also found in the 
Himalayas. 

Geographic Range in the Soviet Union 

Range in the Soviet Union very extensive and comprises about one- 
half of range of species, namely, almost the whole of its northern 
half. It covers the southern region of central Siberia, Trans- Yenisey, 
Siberia, and the Far East excluding the northeastern sector. 

In the southwestern range musk deer occur in the Altai. Extreme 

"More detailed information on morphological characteristics of species can be 
obtained from Flerov (1952) and Tsalkin (1947). 



106 

southwestern habitats of musk deer in the Altai are Kurchum and 
Narym mountain ranges and the upper courses of the Bukhtarma 
(A.M. Nikol'skii, 1883; Kuznetsov, 1948; Antipin, 1941), i.e., proba- 
bly the southern Altai mountain system.' 

Musk deer are dispersed extensively throughout the Altai but it is 
impossible to establish precisely the boundaries of their range there, 
especially in the past. Encounters with musk deer are known in the 
Katunsk range, in the Koksa basin, i.e., Korgonsk (Abai and Kok- 
sinsk mountains), in the upper reaches of Charysh (Ust'-Kan), i.e., 
probably also in Tigerets range, and along the Cherga, i.e., in the 
87 Cherginsk and Seminsk mountains. Farther east the range boundary 
runs to the northern extremity of Lake Telets (Kashchenko, 1899; 
Flerov, 1936; A.M. Nikol'skii, 1899; Yurgenson, 1938; V.G. 
Heptner). There is no positive information about the occurrence of 
musk deer in the western mountains of Ubinsk, Ul'binsk, and Iva- 
novsk Belkakh, but its habitation there, at least in the past, is highly 
probable. The boundary does not run up to the north of the range 
and evidently did not reach Gorno-Altai (Oirot-tury and Ulaly), 
because environmental conditions are unfavorable for the species.'^ 

Farther away the range covers the Abakansk range, at least its 
southern part in the region of Lake Telets (V.G. Heptner) and the 
Kuznetsk Alatau (Flerov, 1935; Yanushevich and Blagoveshchenskii, 
1952). Details of the distribution of musk deer in this zone are not 
known. In the Salair ridge it is evidently absent. East of the Altai, the 
range encompasses the Sayan mountains in the north right up to 
Krasnoyar (Stolba sanctuary) and the Tuvinsk district, and in the 
south includes Tannu-Ola. 

The distribution of musk deer in the rest of the range (greater 
part of Siberia) is poorly known. It is described here in a very general 
manner. The boundaries of the range at several places have been 
outlined only roughly or even tentatively because of inadequate 
information. Yenisey serves as the western boundary of the range 
and musk deer do not go beyond it. Only in a few places such as 
Osinov rapids, near the mouth of the Podkamennaya Tunguska or 
in the region of Alinsk, near the mouth of the Elogui (S.P. Naumov, 

According to some data the range boundary is represented by the northern slope 
of the Narym range(Strautman, 1953). References to the Kurchum range pertain to its 
extreme northeastern section — from Altai village (now Katon-Karagai). Information 
about the propagation of musk deer in Saurakh (Kuznetsov, 1948) is not entirely 
accurate and calls for confirmation. 

References to the occurrence of musk deer in the environs of Biisk (Flerov, 1935) 
are undoubtedly erroneous and probably based on incorrect labeling of an animal in 
the collection. 



107 



40 60 во 70 80 90 100 110 120 130 140 160 tOO 




260 о 260 500 760 km 



100 110 



Fig. 27. Distribution boundaries of musk deer, Moschus moschiferus L. in the Soviet 

Union. Arrow in southwest part of range indicates occurrence in Saurakh; question 

marks in northern part pertain to finds in Khatanga (scale in km). V.G. Heptner. 



1931), is it encountered on the left bank of the Yenisey, right on the 
river itself. The northern limit of distribution of musk deer along 
the Yenisey evidently lies at the mouth of the Lower Tunguska 
(N.P. Naumov, 1934). It probably penetrates slightly more to the 
north. East of the Yenisey, in the north, musk deer have been 
reported at Lake Nekongda (66° N lat.), on the Tembencha River (a 
tributary of the Kochechumo, the northern tributary of the Lower 
Tunguska), in the watershed of the northern tributaries of the 
Kochechumo, Embichima, and Yagtala (N.P. Naumov, 1934), in the 
Syverm mountains at 100° E long. (Middendorff, 1867). The possibil- 
ity is not excluded of musk deer occurring even more northward in 
the Kureika basin and probably in the mountains of Putorana, where 
living conditions for it are evidently favorable at places, but there is 
no positive information to support this assumption. '^ The species 
has been noticed on Lake Yesseiy (Tolstov, 1916) roughly at 69° N 



' There is even a reference that "the latitudes of Turukhansk — m'ddle reaches of 
the Tembencha River — represent the northern limit of transgression of some anim- 
als" (Podarevskii, 1936). This is roughly at 65° N lat. 



108 

lat. This evidently is the northernmost reliable point of encounter of 
the species in the Yenisey-Khatanga region. 

According to some data, musk deer range along the Khatanga, up 
to 70° N lat. (Flerov, 1952), which, considering its distribution in the 
Yenisey region, is dubious and requires confirmation. 

Throughout the immense range stretching from the Khatanga to 
the Sea of Okhotsk the distribution of musk deer and the northern 
boundary of its range are not known for certain and the few scattered 
references available cannot provide a correct picture. Thus, in the 
area between the Khatanga and the Lena musk deer are known from 
the sources of the Ikikit River (Ukykit), the left tributary of the 
Olen'ok (about 69° N lat.; Maak, 1859).^" On the Omolon and in the 
lower reaches of the Yana musk deer have been recorded in the north 
up to 70° N lat., and for the Tulyakh mountains 460 km north of 
Verkhoyansk (Tugarinov, Smirnov and Ivanov, 1934). 

The known distribution of musk deer in the Indigirka basin and 
Kolyma is very vague. ^' Judging from available information it 
can be assumed that musk deer have probably spread along the left 
bank of the Indigirka into the region of its watershed with the Yana 
and roughly up to 70° — up to the northern extremity of the Chers- 
kii range. Along the right bank of the river, between it and the 
Kolyma, it hardly reaches north of 67°30' N lat., which is the north- 
ern end of the Mom' range. Farther away the boundary probably 
encircles the Kolyma lowland from the south and runs into the 
Kolyma somewhere in the region of central Kolyma (Zyryanka) or 
slightly above it. In any case it encompasses the region of the right 
tributary of the Kolyma, i.e., the Korkodonsk (Tugarinov, Smirnov 
and Ivanov, 1934). Running farther through the upper reaches of the 
Omolon it intersects the Kolyma range at 64°, emerging on its 
eastern slope slightly short of the coast of the Penzhina Gulf; with- 

^°Maak (1859) emphasized the absence of musk deer along the Vilyuysk and 
pointed to its habitation only along the Chone, i.e., the right tributary of the Vilyuysk 
(around 62°). In his reference to Ukykit he himself doubted this and suggested that 
they could be assigned to mountain sheep. Probably musk deer were actually present, 
all the more so since in the section on sheep he does not cite these data. Transgres- 
sions of musk deer into Vilyuysk are known (Tugarinov, Smirnov and Ivanov, 1934). 
Data for Vilyuysk and Chone cannot, of course, describe the northern boundary of 
habitation of the species in this part of Siberia. 

^'The statement that musk deer probably reached up to 70° only in Indigirka and 
later descended to 69° (Flerov, 1952) is not supported by accurate data and appears 
incorrect. Even in the map shown in the same work the range runs much farther 
north. Precisely for the same reason the statement by Kolyushev (1936) about the 
habitation of musk deer in Yakutia up to the northern limit of the forest tundra is 
disputed. 



109 

out crossing the Penzhina Gulf, it turns along the eastern slope of 
the Kolyma range south (Belopol'skii, 1932), forming the eastern 
boundary. Thus the northern part of the Kolyma range and the 
mountains along the right bank of the Lower Kolyma and the lower 
reaches of the Omolon, including the Anyui range, evidently do not 
fall within the range of musk deer. 

The eastern boundary of the range of musk deer is formed by the 
mountains on the shore of Sea of Okhotsk reaching up to the sea 
coast (Abramov, 1954).^^ The boundary line due to local terrain 
conditions is fairly complex. Thus, for example, in the plains of the 
Amur estuary region musk deer are absent (Maak, 1859). Farther to 
the south, along Sikhote-Alin, the range evidently runs everywhere 
up to the sea coast or comes close to it. 

The southern boundary of the range within the USSR is repres- 
ented by the state boundary up to which the range reaches almost 
everywhere. Exceptions are the relatively small areas along Amur 
and Ussuri valleys and very insignificant sections in the steppes of 
southeastern Trans-Baikal, in the region of Kyakhta and the lower 
89 reaches of the Dzhida. These areas, in spite of prevailing views, are 
extremely small and no more than "open spaces" in the range, also 
seen in the north.^^ In general, the range everywhere runs beyond the 
state boundary; only along Tannu-Ola (USSR) do the state and 
range boundaries merge in the south. 

Musk deer are found in the northern as well as southern parts of 
Sakhalin, but are absent in the Shantarskiy Islands and Kamchatka. 

Within the range described above, the distribution of musk deer 
is extremely uneven. This is explained by the animal's affinity for 
montane taiga and rocky sections and its pronounced stenotopic 
nature. Therefore the detailed placement of the species as a whole 
in a given region is extremely complex; pockets of habitation of the 
animal alternate with areas of its absence. Regions of habitation can 
be considerable in area. The population of musk deer is also very 
uneven, which makes for a characteristic pattern of disposition of 
animals in individual pockets. 

^^According to some reports (Flerov, 1935 and 1952), based on old data, the 
boundary runs along the coast of Sea of Okhotsk but at some distance from it. 

Musk deer are therefore found even along the Nerchinsk range in the Borzya 
region and are also caught in Byrkov region, slightly east and northeast of Borzya. 
They have also been sighted along the Chikoya, especially in the Kransnochikiisk 
region, and are generally found "everywhere in southeastern Trans-Baikal regions" 
(Podarevskii, 1936). 

Evidently herein lies the reason for the absence of musk deer in the plains and 
lowlands of western Siberia, west of the Yenisey. 



110 

Geographic Range outside the Soviet Union 

In Mongolia musk deer are found in the mountains of Kosogol 
(Hobsogol) region and from there southeast along the Selenga, part 
of the Khangai system to the south roughly up to 45 to 46°, and 
Hentei southwest of Ulan Bator, and the Bogdo-Ula mountains. 
Musk deer are evidently absent in the Mongolian Altai and along 
offshoots of the Great Khingan entering Mongolia in the east. In 
northeastern China musk deer have spread to all the mountain sys- 
tems (Great and Little Khingan, Il'khuri-Alin, east Manchurian 
montane region, and Chanbaishan) and the range thus covers 
almost the entire country except the broad expanses at the center 
(Manchurian plains) and some other areas. 

Farther, the range (reconstructed) covers the Korean Peninsula 
(details not known), eastern parts of Inner Mongolia (absent in the 
west) and northern China (Jehol, Hebei, and Suiyuan') except, of 
course, the desert of Ordos and Alashan, Shansi, and Shensi. The 
western part of the range covers the eastern part of the Nan Shan 
range in the west at least up to Lake Karanor (about 98° E long.), the 
Kukunor region (South Kukunor range and others), eastern edge of 
Tibet (Kama and the region of the tributaries of Huang Ho and 
Yangtze, Mekong and Salween), the southernmost tip of Tibet and 
the Himalayas, and in the west up to Kashmir and Gilgit. The 
southern boundary of the range in China and Indochina is not clear. 
Evidently it includes the northern montane parts of Assam and 
Burma and south China provinces, evidently including Yunnan. 
Available information about the penetration of the species under 
description into south Burma up to Pegu province (in the region 
west of the lower Salween) and in the eastern part of Indochina 
along the mountains up to Vietnam is rather dubious. 

The eastern boundary of the reconstructed range may be tenta- 
tively drawn along the sea coast or close to it along the eastern rim of 
hilly Nan'lin'. At present, since much of eastern China is densely 
populated, musk deer do not reach their natural eastern boundary. 
On the plains between the lower reaches of the Yangtze and Huang 
Ho and slightly north (toward Peking), i.e., in the Great China 
plains, musk deer have evidently been absent from time immemorial. 
(V.H.) 

^^Data for southern Burma and Indochina are fairly old and unconfirmed. It is 
highly possible that confusion between musk deer and Tragulidae existed. 

^^The range of musk deer outside the Soviet Union is based on data from Ban- 
nikov, 1954; Flerov, 1930 and 1952; Tate, 1947; Ellerman and Morrison-Scott, 1951; 
and others. 



Ill 

Geographic Variation 

The intraspecific taxonomy of the species, in spite of several studies 
completed to date, cannot be regarded as adequately understood.^^ 
90 Various researchers acknowledge five to seven geographic races. 
Least understood are forms in the southern parts of the range (Hima- 
layas, southern China, and Tibet). Within the Soviet Union two to 
four subspecies are distinguished (Tsalkin, 1947; Flerov, 1952). 
Worldwide nomenclature is far from consistent. 

In the Soviet Union three subspecies can evidently be distin- 
guished but the differences among them are extremely insignificant, 
the characters overlap, and more precise data are required before 
definitive separation is permissible. 

1. Siberian musk deer, M. m. moschiferus Linn., 1758 (syn. altai- 
cus, sibiricus, and arcticus). Relatively large with maximum skull 
length of 164 mm. Nasals constitute 41% of maximum skull length. 
Interorbital width in most specimens more than width across 
supraorbital processes. Nasals usually form a wedge-shaped figure 
at posterior end. Metacarpals and metatarsals long (former 128 to 
148 mm and latter 172 to 189 mm). Color dark. Hair base white. 

This form is found throughout the Siberian part of the range 
except the Amur and Sakhalin. Outside the Soviet Union it inhabits 
northern Mongolia. The range boundary in relation to the next 
form is quite vague. 

2. Far East musk deer, M. m. parvipes Hoi., 1911 (syn. turovi). 
Generally somewhat smaller than preceding subspecies. Maximum 
skull length up to 157 mm. Nasals relatively short and constitute 
about 38% of maximum skull length. Interorbital width invariably 
more than width across supraorbital processes. Nasals usually do 
not form wedge-shaped figure posteriorly; they are blunt instead. 
Metacarpals and metatarsals somewhat shorter. Color darker than 
that of preceding form, i.e., chocolate- brown. Hair base gray. 

The range covers the Ussuri and Amur. Outside the Soviet 
Union this form is found in Korea and probably adjoining parts of 
northeastern China. This is a very poorly distinguished form and 
reflects a tendency toward reduction in overall body size from north 
to south. Sometimes Amur and Ussuri musk deer are placed under 
the Siberian form (Flerov, 1952). 

3. Sakhalin musk deer, M. m. sachalinensis Flerov, 1929. Rela- 
tively small form. Maximum skull length 151 mm. In skull structure 
similar to Far East musk deer width of interorbital area less than 

^'See the characteristics of the subfamily given below for total number of species. 



112 

width across supraorbital processes, and very rarely almost equal to 
it. Metapodium as in preceding form. 

The range covers Sakhalin Islands. 

Information on the geographic variability of musk deer outside 
the Soviet Union is rather vague and often contradictory. The forms 
described call for a revision. The following forms should be singled 
out for special study: M. m. berezovskii Flerov, 1928 (Szechwan; very 
indistinct form, sometimes designated a special species); M. m. sifa- 
nicus Buchner, 1894 (central China, Kansu, and Szechwan); andM. 
m. chrysogaster Hodgson, 1839 (Himalayas — Kashmir, Nepal, and 
Sikkim; possibly identical with the preceding form). (V.H.) 

Biology 

Population. The distribution of musk deer is characterized by 
uneven dispersal. They are found only in some sections of the habi- 
tat and generally do not form large herds. At places where musk deer 
have long been protected the population density sometimes reaches 
73 to 78 animals per 1,000 hectares of forest area (Boskonsk River 
basin in the Altai preserve, 1940 and Stolba preserve, 1952). At places 
where musk deer are caught in large numbers population density 
does not exceed 5 to 10 animals per 1,000 hectares (Shcherbakov, 
1953; Sliaposhnikov, 1956). Stolba preserve (46,500 hectares) con- 
tains in toto slightly more than 500 musk deer (Dul'keit and Kozlov, 
1958). 

Habitat. In the Soviet Union musk deer mostly inhabit the mid- 
dle altitudes of montane taiga and are not usually found above 1,600 
m. The animals, especially in winter, are attracted to relatively steep 
91 slopes covered with coniferous forest (nut pine, fir, or spruce, and 
rarely larch or Scots pine). Favorite habitats are sections with rock 
outcrops which provide shelter from predators. A distinct preference 
is shown for northern shaded slopes where arboreal lichens abound 
and an ice crust does not form in winter. Musk deer are rarely sight- 
ed in smooth terrains or in places far away from rocks (in the Lena 
basin and upper courses of the Angara this phenomenon occurs 
more often than in other regions); hence their distribution is usually 
mosaic in pattern. In the northern parts of the range they inhabit 
elevations covered with coniferous forest or forest-tundra vegetation. 

In summer musk deer spend at least part of the day in valleys of 
forest rivers, near mountain streams, and along sections with good 
grassy vegetadon where coniferous taiga alternates with mixed 
deciduous forests. They avoid marshy forests. 

Male musk deer, especially in winter, live at a slightly higher 



113 




Fig. 28. Habitat of musk deer in eastern Sayan. Region of Lake Medveshi. July, 
1956. Pfiotograpli by K.T. Yurlov. 

elevation than females; they are absent altogether on gentle slopes 
(Sayans and Altai). 

South of the Soviet Union's boundary musk deer inhabit moun- 
tains covered with deciduous forest and shrubs in sections where 
arboreal lichens are totally absent. Moschus m. berezovskii, which is 
close to the Siberian musk deer (Szechwan and Kansu), is confined to 
the alpine belt, however (Flerov, 1952). 

Food. Over 130 plant species are known to be consumed by musk 
deer (Shcherbakov, 1953). Of these, about 20 constitute the main 
diet. In the Altai musk deer consume not less than 95 plant species, 
of which 9 are particularly important (Shaposhnikov, 1956). In the 
diet of musk deer lichens, primarily arboreal lichens (several species 
of Usnea, Parmelia, and Evernia) and to a lesser extent terrestrial 
bushy lichens {Cladonia and С etr aria), play an important role year- 
round. Lichens are of particular importance in winter, when they 
constitute up to 70% by weight of the stomach contents of musk deer. 
In Stolba preserve the stomach contents of 31 musk deer caught in 
winter contained almost invariably considerable quantities of 
lichens; in summer lichens were found in 50% of the stomach con- 
tents examined and in very small quantities (Shcherbakov, 1953). In 
the Altai lichens predominate in the food of musk deer from Sep- 



Э 



114 

92 tember through May (Shaposhnikov, 1956). In captivity musk deer 
.consume about 0.8 kg lichens per day (A.G. Kostin). 

Musk deer avidly consume arboreal food year-round. This 
includes thin branches and young shoots, coniferous needles and 
bark (mainly in w^inter), leaves, buds, and rarely small rootlets. Arbo- 
real food in w^inter is second in importance to lichens. Musk deer 
consume mountain ash, aspen, maple, willov^^, bird cherry, honey- 
suckle, and other deciduous species; in winter they often nibble nee- 
dles and end branches of fir, sharp-edged yew (in Sikhote-Alin), and 
rarely Scots pine and nut pine (Doppel'mair, 1926; Scherbakov, 
1953; Shaposhnikov, 1956; Yu.A. Salmin). Needles evidently consti- 
tute a compulsory food. In Stolba preserve in winter needles have 
been found in almost all the stomach contents of musk deer and 
usually in considerable quantities. However, in captivity, when 
more satisfying food is available, the animals do not touch needles 
(Shcherbakov, 1953). Musk deer avidly consume in summer leaves of 
common sweetbriar (Stolba preserve) and in winter terminal 
branches and leaves of rhododendron (Sikhote-Alin). In the Irkutsk 
region musk deer consume nuts of Siberian pine (Kopylov, 1950). 

Herbaceous plants serve as the main summer food of musk deer. 
At this time of the year they avidly consume buckwheat, umbellifer- 
ous plants, burnet, spirea, geranium, Gmelin peavine, leaves of bil- 
berry and stone bramble, some grasses, etc.; the total weight of green 
mass in a stomach can weigh up to 2 kg (Shcherbakov, 1953; and 
others). In winter the dry parts of the following plants are also 
consumed fairly regularly but usually in small quantities: aconite, 
cow parsnip, willow herb, ferns, grasses, sedges, roots of some 
plants, and remnant green stalks of bilberry and horsetail (Kopylov, 
1950; Shaposhnikov, 1956; and others). 

In the Altai musk deer consume green moss {Mnium cuspida- 
tum), the roots of arboreal ferns, and pileate mushrooms (birch 
mushroom, pepper mushroom, and others). Sometimes, musk deer 
successfully feed on mushrooms stored by squirrels among twigs 
(Shcherbakov, 1953; Shaposhnikov, 1956). In summer the food of 
musk deer is much diverse than in winter. 

Musk deer obtain conifer needles and arboreal lichens not only 
from trees (up to a height of about 1.5 m) by standing on their hind 
legs, but also from windblown dry branches and sometimes forest 
clearances. Musk deer dig up food from under the snow only when 
the cover is no more than 20 to 30 cm deep (A.G. Kostin and F.D. 
Shaposhnikov). 

In summer musk deer periodically forage in those state farms 






115 



least visited by other ungulates. In captivity they take salt but only a 
small quantity. 

Home range. Musk deer live for most of the year in a home range 
of 200 to 300 hectares, holding onto it steadfastly. Such a home range 
comprises steep rocky "shelters" and more gentle slopes for feeding. 
The size of the home range decreases perceptibly in the second half 
of winter. In summer and during estrus musk deer wander exten- 
sively (Shcherbakov, 1953; Shaposhnikov, 1956; and others). 

In a fir forest in Stolba preserve the average distance of daily 
movement of musk deer in winter was 3.8 km and in a pine grove 
with less abundant lichens, 4.3 km (Shcherbakov, 1953). In the Altai 
the distance covered daily on snow was roughly 4 to 5 km. On being 
chased musk deer usually run round and round in circles within 
their home range (F.D. Shaposhnikov). 

Musk deer usually live singly or in groups of not more than two 
or three animals (adult female with young) and the habitats of this 
species are more isolated than those of other hooved animals. How- 
ever, many musk deer go to distant places to feed on lichens, which 
are more readily accessible; some travel as much as 5 to 7 km 
(Shcherbakov, 1953). 

Daily activity and behavior. Musk deer are active at twilight or at 
night. In summer the animals descend from their lair soon after 
sunset and feed in forest meadows and river valleys and near small 
brooks, reascending 100 to 200 m to their lair by morning. The lairs 




93 Fig. 29. Habitat of musk deer in the Altai sanctuary. Gorge of the Chira River 
visible in foreground. August, 1951. Photograph by F.D. Shaposhnikov. 



116 




Fig. 30. Track of musk deer in deep snow. Upper course of the Kansk River. Eastern 
Sayan. March, 1952. Photograph by K.T. Yurlov. 

93 are set close to rocks and sometimes among rock debris and along 
crests of narrow mountain ranges where, because of sparse forest, the 
wind is strong and hence mosquitoes and midges few. Musk deer lie 
comfortably at the edge of precipices, on the trunks of fallen trees, 
rocks, and so on (F.D. Shaposhnikov; and others). In the mountains 
of Sikhote-Alin musk deer are often troubled by blood-sucking flies 
when in dense grass, or moving near stumps or among brushwood 
left behind after floods. In such circumstances they remain during 
the day in river valleys (Yu.A. Salmin). Inside the lair the animals 
remain hidden and permit close approach. The lair is located in a 
fixed place. 

94 In winter musk deer are active only for part of the night and 
usually lie down one to three times during the hours of darkness. In 
heavy snowfall and inclement weather the animals are less active 
(F.D. Shaposhnikov) and lie down at places which provide protec- 
tion from the wind. 

"Musk deer have long mobile toes ending in sharp and hard 
hooves which are extremely tenacious and provide the animal grip 
not only on the slopes of steep hills, but also on trunks of fallen or 
steeply inclined trees" (Yu.A. Salmin). Their gait resembles that of a 
rabbit. Musk deer invariably use the same pathways and thus form 
permanent tracks. In winter this beaten track facilitates movement 
on loose snow. 



117 

Information about the sense organs of these animals is contradic- 
tory. It is generally acknowledged that vision is keen, hearing good, 
and sense of smell poor. 

Seasonal migrations and transgressions. Musk deer lead a fairly 
well-settled mode of life in most of the range in the Soviet Union. 
Where seasonal migrations do occur, they seldom exceed 1.0 to 5.0 
km. Depending on topographic features, the animals live in winter 
at roughly the same altitudes as in summer, or in some regions 200 
to 300 m (vertical height) below the summer altitudes or, contrarily, 
above them. In winter in mountains of central Sikhote-Alin musk 
deer live in high elevations and along mountain slopes (Yu.A. Sal- 
min). Changeover to winter habitats occurs imperceptibly but usu- 
ally takes place during September. 

In winter musk deer prefer dense boreal forests. The weight dis- 
tribution on the hoof (80 to 120 g per cm^ of resting surface; Dul'- 
keit, 1956) is relatively low and the minimum for ungulates. Their 
dependence in winter on food available above the snow such as 
arboreal lichens and branches of trees and shrubs enables musk deer 
to live even in extremely snowy regions in winter, but they are 
confined to extremely steep slopes. In less snowy sections (with a 
snow cover of not over 50 cm), deer move not only along mountain 
peaks but also along relatively gentle slopes (Dul'keit, 1956). In 
spring, with the appearance of thawed patches, many animals can 
be seen on southern slopes, moving away from them with the 
approach of summer. 

Reproduction. In the Altai, Sayans, Yakutsk Autonomous Soviet 
Socialist Republic, Chita region, and Sikhote-Alin, musk deer usu- 
ally come into estrus in December (in the Altai and Yakutia from the 
end of November) and the period generally extends over three to four 
weeks. However, according to Yu.A Salmin estrus in musk deer in 
central Sikhote-Alin occurs from early November to early December, 
i.e., almost a month earlier than in the Altai. In Stolba preserve 
estrus in musk deer continues throughout the whole of November 
and most of December. Among males spermatogenesis commences 
before rut in September and extends through February. In females 
no distinct pre-estrus stage is evident as the follicles are formed 
throughout the year (Shcherbakov, 1953). 

Unlike other periods of the year in which herd formation is not 
at all characteristic of musk deer, during the period of estrus groups 
of three or four adult animals and in very rare cases even up to five to 
seven (two males and three females, three males and four females, 
etc.) can be seen. The young live independently. A male may mate 
with several females. A single female sought by^everal males is also 



118 

not rare. A more powerful male attempts to chase away the weaker 
but does not generally use his canines for this purpose. Males emit 
a low squeal or "hiss" (Belousov, 1928; Shcherbakov, 1953; Sha- 
poshnikov, 1956). The site of musk deer in estrus is recognizable by 
characteristic "stamping" of the ground and crushed grass. An 
excessive exudation of musk in males in rut evidently exerts a stimu- 
lating influence on females in heat. At the end of the rutting period, 
in early January in the Altai, musk deer groups break up (Shaposh- 
nikov, 1956). The gestation period of musk deer is 185 to 195 days 
and there is no latent stage of embryonic development (Shcher- 
bakov, 1953; Shaposhnikov, 1956). The female usually delivers two, 
rarely one, or even more rarely three fawns. However, in Stolba 
95 preserve, of 13 pregnant musk deer examined, 9 had a single embryo 
and 4 had two each. 

Most females fawn in June and some in May or early July. In the 
Altai isolated births of fawns have been recorded right up to July 12 
(A.G. Kostin). In Sikhote-Alin fawning even commences at the end 
of April (Yu.A. Salmin) and the last fawns are born one month 
earlier than in the Altai. Fawning usually occurs in secluded places 
such as dense shrubs, under low branches of fir, in the proximity of 
fallen trees, etc. In Sikhote-Alin up to one-third of adult females 
remain barren every year (Yu.A. Salmin). 

Musk deer attain sexual maturity at the age of 15 to 17 months 
(Shcherbakov, 1953).'^ 

Growth, development, and molt. Fawns remain helpless for 
quite some time and the mother does not lead them out until the end 
of July or early August. She conceals herself and, in the event of 
danger, tries to drive away the enemy (Shaposhnikov, 1956). Suc- 
kling lasts up to the end of August or early September and the fat 
content of the milk may reach 19%. Fawns lead an independent life 
from autumn or sometimes from spring of the following year 
(Shcherbakov, 1953; Shaposhnikov, 1956; and others). 

Newborns weigh from 300 to 500 g, rarely more. In Stolba pre- 
serve three-month-olds weigh 4.0 to 4.5 kg, which increases to 6.0 to 
7.0 kg by the fifth month. In the Altai in winter young deer weigh 
7.0 to 9.0 kg (Shcherbakov, 1953; Shaposhnikov, 1956; A.G. Kostin; Yu. A. 
Salmin). Adult males and females are almost equal in weight. In 



^*In discussing rut in musk deer in eastern Siberia, the Far East, and Mongolia, 
Cherkasov (1884), Abramov (1954), and other researchers mention vicious fights 
among males during this period which probably is an exaggeration. 

""Flerov (1952) erroneously placed the onset of sexual maturity in musk deer in 
the autumn of their third year. 



119 

Stolba preserve the weight of males averages 13.3 kg (maximum 16.5 
kg) and of females 13.7 kg (maximum 16 kg). Three adult musk deer 
caught in winter in the Altai weighed 10.3 to 16.8 kg. In Sikhote- 
Ahn adult females weigh 15 to 17 kg (Shcherbakov, 1953; Shaposh- 
nikov, 1956; A.G. Kostin, Yu.A. Salmin). 

Juvenile hair molts three to five months after birth and subse- 
quently the young differ very little from adults in color. At about 
one-and-a-half years of age coloration is identical to that of adults 
(Shcherbakov, 1953; Shaposhnikov, 1956). 

Secretion of the musk gland commences in males at the age of 
four to five months (Shcherbakov, 1953) and milk teeth are shed in 
the second year. By the end of that year milk canines have been 
replaced with permanent canines in males; in three-year-old males 
their length often exceeds 60 mm and they become increasingly 
curved. 

Molt in musk deer is a remarkably slow process, continuing from 
mid-February or early March to October or early November. Molt is 
most intense in April and May when the underfur is shed and the 
pelage of the animals extremely sparse. Molt commences from the limbs 
and head, then the abdomen and sides, and last to the croup. New 
hair grows slowly and the back is covered for almost the entire 
summer with a "saddle" of old, highly faded hair, dirty yellow in 
color (Yu.A. Salmin; and others). Among pregnant females the com- 
mencement of molt is somewhat delayed (Shcherbakov, 1953). There 
is no autumn molt among musk deer. 

The life span of musk deer has not been definitively established. 

Enemies, diseases, parasites, mortality, competitors, and popula- 
tion dynamics. Predators from whom musk deer suffer most are lynx 
and wolverine and in the mountains of Sikhote-Alin, yellow- 
throated marten; wolf and the rare tiger cause much less damage. In 
Stolba preserve musk deer remains have been found in 43% of the 
feces of lynx (117 specimens) gathered over a period of three years, 
mainly in winter (Shcherbakov, 1953). In Altai preserve musk deer 
remains were found in four of eight fecal samples of lynx collected 
in July to November. Musk deer badly mauled by wolverines have 
been found time and again in Altai preserve and once the chase of a 
single musk deer by two wolverines was observed (Shaposhnikov, 
1956). 

In the central parts of Sikhote-Alin the yellow-throated marten is 
the main enemy of musk deer, especially in autumn and winter 
when these predators hunt in family groups. The damage caused by 
marten greatly increases in the event of ice crusts, which greatly 
facilitate chase of the quarry. In the winter of 1935 to 1936, on the ice 



120 

96 of the Arma River basin, over a distance of 200 km the remains of 26 
bodies of musk deer torn to pieces by martens were found (Yu.A. 
Salmin). In spring and summer young deer are particular victims of 
martens. The population dynamics of marten in some years depends 
greatly on the population of musk deer (Bromlei, 1953 and 1956). 

Only in rare cases do musk deer become the quarry of brown 
bear. In the Altai, of the 180 samples of feces of this predator exam- 
ined, the pelage of young musk deer was found in two (Shaposh- 
nikov, 1956). Kaplanov (1948) found the remains of musk deer torn 
to bits by a litter of tigers in Sikhote-Alin mountains. In very rare 
cases sable, red fox, horned owl, and golden eagle attack musk deer 
(Kozhanchikov and Kozanchikov, 1924; S. Naumov, 1931; Shaposh- 
nikov, 1956). Remains of musk deer killed by martens are often 
consumed by Siberian weasels (Bromlei, 1953). 

In rescuing itself from a pursuer a musk deer will attempt to 
climb onto a rock, into a "shelter," or finding neither run in circles. 
A marten tries to cut across the deer's arc or chase it onto ice. Musk 
deer can run exceptionally fast only for 200 to 300 m and thereafter 
tire. Dogs and wolves soon outstrip musk deer over relatively level 
surfaces (Cherkasov, 1884; Bromlei, 1953 and 1956). Evidently preda- 
tors constitute one of the reasons for the absence of musk deer on 
relatively gentle slopes, away from rocks. 

The diseases musk deer suffer from have not been studied much. 
In autumn of 1919 in the mountains near Lake Telets dozens of 
bodies of musk deer were found, which had died from some 
unknown disease. According to old hunters such an epizootic has 
occurred only once (A.G. Kostin). In zoological parks mortality of 
musk deer due to necrobacillosis has been recorded. A musk deer 
which survived in a zoological park for three years (a record!) died of 
tuberculosis (Kadenatsii, 1958). In the Soviet Union, under condi- 
tions of zoological parks, 11 species of helminths have been identi- 




Fig. 31. Winter feces of musk deer (magnified). Primor'e territory (from 
Formozov, 1952). 



121 

fied for musk deer — 2 species of cestodes and 9 species of nematodes. 
However, some of tfiese are parasites transmitted by otfier animals. 
Under natural conditions musk deer serve as hosts most often for 
Setaria cabargi and Pneumocaulus kadenazii (Kadenatsii, 1958). 
Parasites found in musk deer in the Altai include Cysticercus taenui- 
collis and Dyctyocaulon among others (Shaposhnikov, 1956), in 
Kazakhstan — Nematodirus (Sokolova and Boev, 1950), and in 
Sikhote-Alin — S. cabargi. Taenia hydatigina, гпб. Monieza expansa 
(Kadenatsii, 1958). 

Musk deer are greatly troubled by mosquitoes, midges, ticks, and 
other insects. More than a hundred ticks were removed from the 
body of a musk deer on one occasion (Stolba preserve; Shcherbakov, 
1953). Almost 2,000 larvae of P avlovskiomyia mexpectata, a parasite 
of the dipteran group, were found under the skin of a Ussurian 
musk deer. Such musk deer become greatly emaciated and in 
summer their back, neck, shoulders, and croup are completely ulcer- 
ated (Abramov, 1954). 

Musk deer have almost no competitors. In winter in the Altai 
musk deer quite often consume lichens fallen from the branches of 
trees shaken by the antlers of maral or broken by bear (Shaposh- 
nikov, 1956). 

Information on the population of musk deer is inadequate. In 
many regions of the Soviet Union, particularly in Primor'e, musk 
deer increased markedly immediately after their hunting was res- 
tricted in the 1930's and destructive methods of catching them prohi- 
bited. Among captured musk deer the sex ratio varies from 4:6 to 
6 : 4 in favor of males. 

Field characteristics. Musk deer are the smallest deer inhabiting 
the USSR and have a characteristic color that makes it impossible to 
mistake them for other ungulates. Seasonal differences in their fur 
color are insignificant. Neither males nor females bear antlers. In 
the field the color of their coat makes sighting difficult. The canines 
97 project beyond the upper lip in males three or four years old or more 
but never in females. The usual sound emitted is a faint hiss. The 
hind legs are much longer than the forelegs and during rapid jumps 
extend beyond them, due to which the hoofprints resemble those of 
hares. On level ground the deer can jump a distance of up to 4.0 m; 
more often, however, this distance is not more than 2.5 m. A fright- 
ened animal may jump up to 4.7 to 5.0 m. On snow, walking prints 
measure 25 to 30 cm between steps (Dul'keit, 1956). 

The toes are capable of considerable movement and hence hoof- 
print dimensions vary widely, depending on terrain and pace. In 
general they are smaller than those of other taiga ungulates but 



122 

similar to those of mountain antelopes* (Formozov, 1952). In winter 
it is very easy to identify hoofprints as male or female from the form 
and position of the "urine spot" on the snow. In addition to tracks, 
hoofprints, and lairs, the presence of musk deer is also revealed by a 
unique feature — multiple feces deposited in one place. Fecal pellets 
are very small (0.7 to 0.8 cm x 0.35 to 0.40 cm) (Formozov, 1952); those 
of males have a pungent "goaty" smell (Shaposhnikov, 1956). (A.N.) 

Economic Importance 

Musk deer are caught mainly for musk ("musk deer perfume"), 
present only in males. Musk is secreted by a saccate gland located 
between the sex organs and the naval. In the middle of the last 
century several thousands of musk sacs of musk deer were exported 
from Russia to China through Kyakhta every year (maximum in 
1855 — 81,200 sacs), but later the animal stocks evidently diminished 
gready. In the hunting season of 1927 to 1928 in the USSR 5,089 
musk sacs were collected by a centralized organization (Yakutia — 
114, Far East — 3,136, and Buryat-Mongol Autonomous Soviet 
Socialist Republic and eastern Siberia — 1,839). Another 193 sacs 
were collected in the present Tuva autonomous region (Nasimovich 
and Rudanovskii, 1935). Today several thousand musk sacs are col- 
lected annually. Musk deer stocks are evidently not fully utilized at 
some places, while in other regions the population of this species is 
low. 

In the past musk was used in medicine in Europe and the East 
and musk imported for this purpose from Siberia for a long time. 
Musk as a natural perfume base in preparing high quality scents 
was discovered later. ^° Musk owes its shaф smell to muskone, a 
ketone-type compound present to the extent of 1 to 2%. Musk is also 
used in the preparation of scented artificial baits in catching preda- 
tory animals. The weight of a musk sac with contents in an adult 
deer ranges from 20 to 52 g; the actual musk weights 15 to 30 g, on 
the average about 20 g. In immature animals the musk content is 
much less. In the period of rut part of the musk is used by the animal 
and its reserves restored within two or three months (Shcherbakov, 
1953). 

The hide of musk deer is not durable as it has brittle hairs which 
deteriorate rapidly. Local people make leggings, mittens, and sleep- 
ing bags out of deer skins as well as use them for rugs. A thin but 

*Reference not clear; possibly goral — Sci. Ed. 

™A satisfactory substitute for musk is the synthetic preparation "tibetolite". 



123 

strong chamois is made from shaven skins and used for pillow cases, 
gloves, etc. The hair is used as mattress stuffing (Abramov, 1954). 

The meat of musk deer is lean and poor in quality; it is most 
often used as bait for catching sable and other fur- bearing animals 
or dried and stored for dog food. The canines are converted into 
various articles. 

In the past musk deer were caught mainly in the countryside 
(enclosed and open pastures) by setting nooses or digging pits in 
their tracks. Nooses set up in tracks were the most common method. 
The end of the noose was tied to a stick which the musk deer could 
drag for some time, until it caught fast in something, after which 
98 the loop tightened on the animal (Petri, 1930). A large number of 
animals were caught by trapping (with nooses and sticks) and plac- 
ing other mechanical catching devices on their tracks. In summer 
hunting used to be done by shooting; the hunter used a birch pipe to 
imitate the call of a female deer and thus attract adult males and 
females. These methods of catching musk deer are now banned. The 
animals are shot from a concealed position, or caught by chasing, or 
bagged by some other method. Sometimes dogs are used to chase 
musk deer toward the hunter or for keeping the animal at bay. Deer 
are shot with bullets and buckshot. Hunting usually occurs from 
September 1 through March 1 (in some parts for a much shorter 
period) and generally requires a license. In 1954 hunting for musk 
deer was prohibited in northern Kazakhstan,, the Altai, Krasnoyar 
region, and Sakhalin. 

Some methods of catching musk deer live are' also known: nets 
(successfully used in the Altai), nooses set at the end of a pole (to 
catch the animal emerging from its "shelter"), and other methods 
makes a preferential catch of adult males possible. 

From the end of the 1930's to the early 1940's in Altai preserve 
experiments were conducted for raising musk deer in captivity (A.G. 
Kostin).^' Later, similar work was undertaken in Stolba preserve 
where about 12 animals were held in a nursery from 1951 to 1954. 
Offspring were raised successfully from two feinale deer. Experi- 
ments on live extraction of musk have not produced promising 
results (Shcherbakov, 1953; Dul'keit and Kozlov, 1958). 

Musk deer are relatively rare in zoological parks. Methods of 
holding this animal in captivity have been studied little and captive 
animals usually die soon. In 1954 a small group of musk deer were 

According to Solov'ev (1922) experiments on raising musk deer in captivity for 
obtaining musk in Ussuri region were conducted even before World War I. The 
correctness of this information is dubious. ^ 



124 

transferred from Stolba preserve to Denezhkinsk Kamensk preserve 
in the Urals and set free for purposes of acclimatization. Tracks and 
deer were sighted there for at least a year but the experiment of 
acclimatization eventually proved positively unsuccessful. (A.N.) 

TRUE OR PLESIOMETACARPAL DEER 

Subfamily Cervinae Baird, 1857 

Moderate to large-sized deer. 

Toes four. Lateral toes usually relatively poorly developed but 
with phalanges. Only proximal part (plesiometacarpal extremities) 
of metacarpal bones retained; cuneiform distinct from naviculo- 
cuboid. 

Facial part of lacrimal with distinct depression for preorbital 
gland. Orbital region of lacrimal with two tear ducts. Nasal process 
(pr. nasalis) of premaxillae does not reach frontal. Ethmoid pit 
large. Vomer relatively short and low at rear and does not separate 
choanae (posterior nares). Because of intense development of fron- 
tals, extending markedly backward, parietals greatly reduced. Apo- 
physes of frontals, serving as base for antlers, relatively short and do 
not form ridges on frontals. Antlers invariably present (only in 
males) and relatively large. Fully developed antlers have more than 
two tines; first tine (brow tine) invariably developed but unbranched. 
Sometimes small flattened "spatula" occurs at tip of antlers. 

Upper canine'small, rudimentary, or absent. Dental formula: 

i |, с ^, pm |, m|- = 32 to 34. 

General build extremely slender and hind legs only insignifi- 
cantly longer than forelegs. Tail moderately long or greatly reduced. 
99 Naked portion at end of muzzle large. Toe cushions developed rela- 
tively poorly and occupy only roughly two-thirds of proximal part 
of hooves, projecting gradually forward into horny substance of 
hoof. Preorbital gland well developed. Metapodial glands present; 
sometimes significant gland clusters occur on tail; no musk or pre- 
putial gland present on abdomen nor any gland on outer surface of 
thigh. Glans penis without filiform protuberance. Preputial part of 
penis attached to abdominal skin. Gall bladder absent. Teats four. 

Representatives of subfamily Cervinae are forest-steppe, forest, or 
montane forest forms, also found in swampy regions and in sub- 
alpine zones. 



125 

Cervinae represent a group of Old World deer. Only one species 
(Cervus elaphus) occurs in the New World and in North America is 
a relatively recent immigrant. In the Old World Cervinae are distrib- 
uted in the south up to the limits of distribution of the family, and 
in the north roughly up to 55 to 60° N lat. (central parts of Norway 
and upper course of the Lena). In the New World they are found in 
North America roughly up to 60° N lat. on the north and 35° N lat. 
on the south (for details, see description of genus Cervus and red 
deer, Cervus elaphus, which follow). 

Subfamily Cervinae in the area under consideration (described 
later) represent a well-defined, relatively monotypic group. These 
deer were placed in different taxonomic divisions by researchers for 
quite some time. Often Elaphodus and Muntjacus, and recently 
CapreoLus and A Ices, have been included in this plesiometacarpal 
group (Flerov, 1952). Group Cervinae may be considered a fairly 
natural group although its separation from Odocoileinae is based 
on relatively insignificant characters. Differences from Muntjacinae 
and Hydropotinae are more significant. 

Cervinae are known from the Lower Pliocene of Europe and 
Asia; their occurrence in the Pliocene of North America has not been 
proved (relationship of the Upper Pliocene Procoileus, known from 
North America, to Cervinae not yet clear). Migration of Cervinae to 
North America probably took place in the Pliocene. 

This subfamily, until very recently, was in an altogether un- 
satisfactory state from a taxonomic point of view. Concepts regard- 
ing the number, extent, and grouping of species were far from clear. 
First of all the number of species is very large and, secondly, special 
generic or subgeneric names have been assigned to almost all the 
well-defined species. Even forms which have been universally 
acknowledged as subspecies have been placed in different genera and 
subgenera. Various researchers hold various opinions. This is often 
true even now^^ but recent studies (Heptner and Tsalkin, 1947; Eller- 
man and Morrison-Scott, 1951; Flerov, 1952; Haltenorth andTrenze, 
1956) have clarified the general picture and differences of opinion 
pertain to details, mainly number and extent of genera and subgen- 
era. The number of species may be taken as established and changes 
are possible only in relation to some forms inhabiting southeastern 
Asia, primarily for downgrading them to the rank of subspecies. 



It is significant that the Siamese Cervus schomburgki (now probably totally 
extinct) has been regarded by some as a special subgenus (Thaocewus) of genus 
Cervus (Ellerman and Morrison-Scott, 1951). Others place it in the subgenus of 
barasingha (C. duvauceli; Haltenorth and Trenze, 1957). ^ 



126 

The subfamily consists of eight genera (about 20% of all genera 
in the family) of which six are extinct and two extant" (about 16% of 
extant genera in the family). Extant genera are red deer (Cervus) and 
the Chinese or Pere David deer (Elaphurus). The remaining names 
proposed for species of the subfamily signify nothing more than 
their belonging to subgenera of genus Cervus. The total number of 
species is 15 or 16; genus Elaphurus is monotypic and the rest 
belong to genus Cervus. This number constitutes about 44% of the 
total number of species in the family.^'* 
100 In the Soviet Union only one genus is found, red deer Cervus 
(50% of genera of the subfamily) with an autochthonous species 
(about 14% of the number of species in the subfamily). ^^ Species of 
the subfamily comprise about 0.7% of the number of species present 
in Russian fauna. 

In the Soviet Union Cervinae are found in the western part of the 
forest zone (western part of the region of broad-leaved forests), in the 
Crimea and the Caucasus, along the southern rim of the taiga, in 
Amur-Ussuri forests, and at places in Central Asia. In the last cen- 
tury, as a result of extinction, the geographic range shrunk at some 
places; as a result of acclimatization experiments from 1930 new sites 
of habitation have been created, partly within the old range and 
partly beyond it. Thanks to conservation measures implemented in 
the post-Revolution period, the deer population has increased in 
several places. 

Cervinae are game and table animals and often objects of sport. 
In some parts of the range deer hunting is forbiddeen. (V.H.) 

Genus of True Deer 

Genus Cervus Linnaeus, 1758 

1758. Cervus Linnaeus. Systema Naturae, 10th ed., vol. 1, p. 66. Cervus 
elaphus Linn. 1827. Elaphus. Hamilton Smith. Griffith's Cuvier 
Anim. Kingdom, vol. 5, p. 307. Cervus elaphus Linn. 

^^The number of fossil genera according to Simpson (1945); however he considers 
four extant. 

^"Pere David deer (mi-lu in Chinese) were found in the northeastern part of China 
proper (Hubeh, Shantung, and Henan) in prehistoric times and confined to flat 
marshy places. This deer became extinct (partly died as a result of the drying up of 
marshes) and before the commencement of the nineteenth century survived only in 
the palace gardens near Peking, from where it entered Europe. At present there are a 
few hundred living in parks in England (Woburn), Australia, and in zoological 
gardens. 

'^Fallow deer (C. dama), as an acclimatized species, have not been taken into 
consideration. 



127 

18S8. Harana. Hodgson. Ann. Mag. Nat. Hist., vol. 1, p. 154. Cervus 

wallichi Cuv. 
IMl.Pseudocervus. Hodgson. Ann. Mag. Nat. Hist., vol. 1, p. 154. 

Cervus wallichi Cuv. 
1846. Strongylocerus. Owen. Brit. Fossil. Mamm. Birds, p. 470. 

Cervus elaphus Linn. 
1870. 5г^й. Sclater. P.Z.S.L., p. 115. Cervus sika = C. nippon Temm. 
1872. Pseudaxis. Gray. Cat. Ruminants Brit. Mus., p. 70. Cervus 

taiouanus Blyth. (subspecies C. nippon Temm). 
1874. Elaphoceros. Fitzinger. Sitzungsber. d. Akad. d. Wissensch. 

Wien, vol. 68, p. 347. Cervus sika Temm. = C. nippon Temm. 
1898. Sica. Trouessart. Cat. Mammalium, p. 878 (substitute for Sika 

Sclater). 

1898. Sikaillus. Heude. Mem. Hist. Nat. Emp. Chin., vol. 4, p. 98. 
Cervus sika Temm. = C. nippon Temm. 

1899. Eucervus. Acloque. Faune de France, Mamm., p. 71. Cervus 
elaphus Linn.^^ 

Cervus are plesiometacarpal deer of large, moderate, and small 

37 

Size. 

Skull relatively elongated and narrow, but fairly broad and short 
in some. Orbits relatively small, but fairly large in some. 

Upper canine usually present in male, although very small, but 
absent in some; rare in female. Dental formula: 

i §- , с ^-^, pm I , m I = 32 to 34. 
6 Jo 3 

Milk teeth moderate in height, sometimes with a fairly high and 
broad crown, but in some animals brachyodont. Sometimes an addi- 
tional cusp occurs on upper molars on the inside. 

Antlers vary in size from moderate to very large and are set some- 
times at an acute angle; in some individuals highly inclined forward 
or directed sideways. In number, size, and form of tines antlers 
extremely diverse. Invariably first tine (brow tine) present and at 
least two others. Number of tines on antlers usually 10 or a few 
101 more. Antlers invariably circular or oval in cross section; in one 
case (C. dama) tip flattened into "spatula". In most (except C. dama 
and the group Rucervus) tines set forward from main beam. 

Synonyms are given only for species inhabiting the Soviet Union. 

"Height at shoulders ranges from 150 cm (maral) to 65 to 70 cm (Philippine deer, 
C. philippensis alfredi). 

^4n С schomburgki. Individuals with 20 to 30 tines, found in some forms of red 
deer (C. elaphus), constitute an extremely rare exception. 



128 

Tail relatively short, of moderate length, or long (length may 
exceed ear length). Hind hooves covered by skin fold almost 
throughout their length. Preorbital gland large; it bulges in some 
individuals and is small in others. Metatarsal glands may be absent; 
if present, well developed, small, and either glabrous or with tuft of 
long hair. Short mane usually present on neck.^^ 

Coloration of young deer spotted or uniform; in adults uniform 
or spotted in winter and summer. Speculum on posterior surface of 
rump varies in size and with or without dark border; sometimes 
totally absent. Black color along middle of tail absent with some 
exceptions. Female somewhat smaller and more slender than male. 

Cervus undergo two molts every year. 

There are forest, montane forest, and forest-steppe forms, but 
partly inhabit swamp forests. 

The geographic range of the genus (reconstructed) is divided into 
two parts — American and Old World. In Africa the range covered 
the northwest, including Morocco, and probably extended up to the 
Spanish Sahara (northern parts of Rio de Oro).^° In western Europe 
the range encompassed the western parts of England and Ireland, in 
the south the Mediterranean countries except Crete and Cyprus 
(Corsica and Sardinia fall in the range), and in the north the south- 
ern half of Norway and Sweden. In eastern Europe the northern 
boundary of the range traversed evidently along a line from Gulf of 
Riga, south of Moscow to the southern part of the central Urals, and 
in Asia from the central Urals to the northern banks of Baikal, 
central Lena and toward the Amur estuary. In the east the range 
encompassed the islands of Japan (including Ryukyu), Taiwan, 
Hainan, the Philippines (absent in Talaud group), Moluccan 
Islands, particularly Batian nd Dzhilolo (Halmahera; possibly 
acclimatized there from antiquity), Seram Islands, Timor, and 
Kambing adjoining Timor from the north (absent on Wetar), and 
the entire island complex north and west of the above places. On the 
Asian mainland the range of the genus covered Indochina, Malacca, 
Ceylon, India, Iran, Iraq, Asia Minor, Syrian region of the United 



In view of the diversity of form in the genus and the abundance of species, 
morphological characters are mainly those of Russian species. 

Information relating to distribution up to Senegal-Gambia inclusive (Lydekker, 
1915) is dubious. Deer in small numbers are found at present only in the forest region 
at the border of Algeria and Tunisia. Information about Egypt is contradictory: 
according to some data fallow deer were acclimatized there even in antiquity, but 
according to other sources the animal was autochthonous there. Occurrence of Dama 
schaejeri, described recently for Egypt, is dubious. 



130 

Arab Republic, Palestine, and the entire area up to the northern 
boundary in Siberia referred to above. In the New World the range 
covered North America roughly between 60° and 35° N lat. (for 
more details, see the section on red deer). From the range character- 
istics the genus should be regarded as Euro-Asiatic, associated lar- 
gely with Asia, mostly its southern and southeastern parts. 

Different forms of the genus are widely dispersed. Sambar were 
acclimatized in Bonin Island and Guam and then imported by local 
inhabitants even in the pre-European period into New Guinea, 
where they were originally absent. In the last decade deer (American 
and European forms of red deer, sambar, and spotted deer) have 
been acclimatized in New Zealand and South and North America. At 
several places in Europe deer had become extinct and their popula- 
tion was artificially restored later. 

Genus Cervus in its characters and number of species occupies a 
central position in the subfamily. Genus Elaphurus (Pere David 
deer) is more isolated and exhibits no close generic association with 
genus Cervus. 

Genus Cervus is known from the Middle Pliocene (or upper 
period of the Lower Pliocene; Flerov, 1950) of Asia. It evidently 
103 prospered in southern Asia where, even at present, maximum diver- 
sity of species of the deer group Axis, known from the Lower Plio- 
cene, is seen. The oldest forms of the genus belong to the group 
sambar (Rusa). In Europe representatives of the genus partly belong 
to its more primitive branches (type of sambar and sika deer, C. 
nippon), known from the Upper Pliocene. Deer migrated from Asia 
to North America only in the Pleistocene, where one of the Euro- 
Asiatic species (C. elaphus) has reached. In origin and in present 
distribution the genus should be considered predominantly south 
Asiatic. 

The number of fossil species of the genus is quite large but all of 
them fall into extant groups within the genus. Some recent species 
can be traced to the Upper Pliocene in a relatively less altered state. 

The taxonomy of the genus until recently was far from satisfac- 
tory. This is primarily true of the number of species, considered 
large even during the period when narrow concepts about species 
prevailed. The grouping of forms proposed and followed by various 
researchers was also extremely diverse and on the whole not really 
acceptable. 

The genus consists of six subgenera: Axis (Indian spotted and 
"porcine" deer) with four species {axis, porcinus, kuhli, and kalami- 
anensis); Rusa (sambars of southern Asia and Australia) with four 
species {unicolor, equinus, timorensis, and philippinus); Przevals- 



131 

kium (Tibetan white-breasted* deer) with one species (albirostris); 
Dama (fallow) with one species (dama — D. mesopotamica, a subspe- 
cies of С dama); Rucervus (barasingha or swamp deer of Indochina 
and India) with two or three species {eldi, duvauceli, and schom- 
burgki, the last evidently only a subspecies of the second); and Cer- 
vus s. str. (red deer) with two species {elaphus and nippon). Other 
subgenera have also been isolated {Panolia, Sika, Taocervus, and 
Hyelaphus), but such a division is superfluous (proximate species in 
such a division often fall in different subgenera). During the period 
of extreme division even well-defined races of the same species (C. 
elaphus) were placed in different subgenera. It is highly probable 
that subgenus Przevalskium, and probably also Rucervus, should be 
placed among Cervus s. str. and only four subgenera recognized. 

In toto the genus comprises 13 or 14 species or about 93% of 
species of the subfamily. Of these, 12 or 13 are purely Asiatic and one 
(C. elaphus) occurs mostly in Asia, but is also found in American, 
European, and African parts of the range of the genus. 

These are game and table animals caught for their meat, hide, 
and antlers in velvet. At places they represent the main object of 
regulated hunting. Hunting of species which are few in number is 
prohibited. C. schomburgki is extinct (Siam). 

Two species are found in Russian fauna — spotted [sika] deer (C. 
nippon Temm., 1937) and red deer (C. elaphus Linn., 1758) — 
representing 14 or 15% of species of the genus and constituting 0.7% 
of species of Russian fauna (mammals). 

These two species are inhabitants of deciduous and mixed forests 
in the west European part of the USSR, the Caucasus, southern 
fringe of the taiga, mountain forests of southern Siberia, and central 
Asian tugais [vegetation-covered bottomlands]. 

These are mainly game and table animals, the hunting of which . 
is often banned. At places they are raised in a semidomesticated state 
for purposes of obtaining antlers in velvet. (V.H.) 

3. SPOTTED [SIKA] DEER 

Cervus (Cervus) nippon Temmink, 1838 

1838. Cervus nippon. Temmink. Coup d'oeilsur la fauna des lies de 
la Sonde et de I'Empire de Japon, p. 22. Japan. 

*Error in Russian original; should read white-lipped. 

""in Soviet literature the following names are common for these deer according to 
sex and age: male — "olen' " [buck], female — "olenukha" [doe], and young two-year- 
old male with simple unbranched antlers ("sack," "saenysh," or "shil'nik"). 



132 

1845. Cervus sika. Temmink. Fauna japonica, Mamm., p. 54, fig. 17. 

Japan. 
\9>&\. Cervus pseudaxis. Gray. Proc. Zool. Soc, London, p. 236. 
I86i. Cervus hortulorum. Swinhoe. Ibid., p. 169. Park of Summer 

Palace near Peking. True native place of these animals is 

Manchuria. 
\д)Ь'\. Cervus mantchuricus. Swinhoe. Ibid., p. 169. 
1876. Cervus dybowskii. Taczanovski. Ibid., p. 123. Southern part of 

Ussuri region. 
1899. C^ri;M5 mantschuricus major. Noack. Humboldt, vol. 3, p. 9. 

Suifun, Ussuri region (?). 
1910. Cervus hortulorum typicus. Ward. Rec. Big Game, 6th ed., p. 

52. Manchuria.''^ 

Diagnosis 

Summer fur of animals of all ages spotted (in winter the spots are 
poorly visible or not developed at all). Speculum very small and does 
not extend above caudal base on croup. Antlers usually with no 
more than four tines each. Tail long, almost equal or equal to 
length of ear. Height at withers not more than 112 cm. (V.H.) 

Description 

Animals of moderate size, very slender and elegant, and the most 
slender of all Russian deer. Neck long, almost vertical, and head 
held high. Croup slightly higher than withers. Ears relatively short, 
proportional. Tail relatively long and equal or almost equal to ear 
length. 

Antlers slender, relatively light, small, set very straight, and tips 
fairly close-set. Beam circular in cross section. Tines on full antler 
four: one brow (bez tine absent), one central, and two terminal set in 
sagittal plane. Crown usually not developed but sometimes third 
terminal tine seen exceptionally in old and particularly powerful 



Apart from these actual synonyms for Ussuri forms Soviet literature also uses 
the names C. dama and C. axis for sika deer. In actual fact C. axis refers to the Indian 
species. 



133 

animals, forming an incipient crown/^ Sometimes rudimentary bez 
tine present, i.e., six tines. The latter, however, is an abnormality 
and the antlers under these conditions generally exhibit features of 
irregular development. In very old animals, especially those with 
additional tines, the clearance between antlers (distance between 
terminal parts of the beam) is more, even extraordinarily — almost as 
105 much as in European deer. Color of antlers dark brown; tips of tines 
in well-developed antlers smooth and yellowish-white. 

Individual color variability is fairly high with regard to general 
color of fur and number, disposition, and color of spots. The main 
shade of fur in type forms is rusty. Along the spine the fur is some- 
what darker, becoming lighter (ocherous and dirty white) along the 
bottom on the sides and on the back. The rusty color extends onto 
the legs also where, however, it is pale. The neck and underbelly are 
pale yellow or dirty gray and the head pale brown. A dark band runs 
from the occiput along the neck and back, disappears on the sacrum, 
appears again at the caudal base, and extends right up to its end. A 
pale yellow band forks off the dark band of the tail on each side. The 
tail at the base is bare. Downward and to the side, from the caudal 
base to the rear of the thigh, a triangular speculum of white hair 
forms. It points downward and merges with the white color of the 
inside of the thighs and inguinal region. The speculum is extremely 
small and bounded on top by a dark band. The hair on the speculum 
is somewhat longer than on the trunk, reaching 2.5 to 3.0 cm in 
length*; this hair stands on end when the animal is frightened. 

Against the main rusty background on the trunk scattered, pure 
white, distinct spots occur. They are smaller on the back and gradu- 
ally enlarge toward the abdomen. On the sides of the body the spots 
fuse here and there and form white longitudinal bands of up to 10 
cm in length. The arrangement of spots shows distinct individual 
variations but two discontinuous rows are generally seen on both 
sides of the spine and orie each on the sides. The coat of males differs 
from that of females in that the hair on the head is dark gray, with 
brown and longer hair on the neck and throat. 

""'ЗисЬ antlers are not seen at present (Bromlei, 1956). Formerly, when there were 
many animals, deer with antlers with five tines were evidently encountered more 
often in taiga (Menard, 1930). The report about 15-year-old animals in Hagenbeck 
Zoological Garden (brought while young from Sidema Park near Vladivostok) with 
10 tines and antlers not inferior in power and beauty to those of German deer 
(Menard, 1930) appears dubious. In any case this is an abnormal, exceptional 
phenomenon. 

♦Some confusion with regard to length of hair on the speculum occurs in the 
text. Contradiction is apparent two paragraphs later — General Editor. 



134 




104 Fig. 33. Antlers of wild sika deer (C. n. hortulorum) from Ussuri region 3.5, 
over 4, and 10 to 12 years old. 

The general hue of the winter coat of females is grayish-brown, 
the head light dirty gray, and the inside of the ear pale gray. Along 
the back runs a faintly discernible dark brown band which enlarges 
toward the rear. The central portion of the tail toward the base is 
dark brown and white along the sides. On the hind quarters occurs a 
small white speculum bounded on the upper side by a dark brown 
band which fuses with the band on the back. The legs are grayish- 
brown, darker in front and much lighter behind. The inguinal 
region and upper inner surface of the legs are whitish, and the lower 
part of the chest dirty brown. Light-colored spots occur on the back, 
upper portions of the sides, thighs, and shoulders. Their number in 
winter is significantly less than in summer and the outlines are not 
sharp. Spots are not seen in all adult animals and, moreover, by 
spring, because of wear of the fur often disappear even in those 
individuals in which they were apparent from autumn. Hair length 
on the trunk, 5 to 7 cm. 

Long dark hair grows in the form of a small mane on the neck 
and nape in the winter coat of males; this is not so in females. Males 
have much darker hair on the chest and along the middle of the belly 
(in the preputial region). In some animals the entire fur is uniformly 
rusty in coloration but darkening (to deep brown) of the neck hair 
has also been recorded."^ 

''''This description comes from Bromlei (1956) and pertains to wild animals (not 
held in zoological gardens). 



135 

Young animals in their first summer coat are very similar to 
adults but their spots are larger and also occur along the middle of 
the back and on the sides. In their first winter coat spots are often 
present and usually well defined. 

Preorbital glands are well developed. Tarsal glands are oval, 
elongated, and covered with bristly hair which is much longer than 
the surrounding hair and much lighter in color. 

The skull is typical of the genus and subgenus Cervus and very 
similar to that of red deer ( С e/ap/zu5). 'Differences in sika deer, apart 
from overall small size, include considerable enlargement of the 
cavity of the typanic bullae, which is convex and rounded, without 
ridges and crests on the surface. The corresponding parts in red deer 
are not enlarged but narrow, and the bullae relatively slightly con- 
vex and angular (Flerov, 1952). 
106 Males and females attain full growth and weight by the third 
year. The size of animals three years old and older are as follows: 

Males (five specimens): body length from tip of muzzle to anus 
168 to 180 cm (M 173); height at withers 104 to 112 cm (M 109); chest 
circumference 114 to 130 cm (M 122); distance of hind hoof from 
hock 47 to 52 cm (M 50); tail length without end hair 17 to 19 cm (M 
18); length of ears 17 to 20 cm (M 18.6); weight 104 to 131 kg (M 
117.4). 

Females (12 specimens): body length 149 to 174 cm (M 162); 
height at withers 87 to 98 cm (M 94); chest circumference 96 to 112 
cm (M 104); length of hind sole 44 to 49 cm (M 46); tail length 
without end hair 14 to 17 cm (M 16.3); length of ears 16 to 18 cm (M 
17); weight 60 to 83.7 kg (M 73.1). 

The length of antlers of adult animals (with four tines) varies 
from 65 to 79 cm and weight from 810 to 1,260 g (eight specimens). 
An exceptional antler of a very old animal with six tines weighed 
1,887 g. Antlers of wild deer are known to reach 90 to 93 cm in 
length (Menard, 1930). The maximum skull length is 320 cm. 

In captivity sika deer interbreed readily with Manchurian wapiti. 
Formerly, when sika deer were abundant in the taiga, such hybrids 
were found so frequently in nature that they had a special Chinese 
name. Hybrids are intermediate in size between parent species, but 

''^All the morphometric data pertain to wild deer and are taken from Bromlei 
(1956). 

Sika deer are called "hua lu" ("flower deer"), Manchurian wapiti "ma-Iu," and 
their hybrids "chin-da-guiza". Antlers in velvet also differ in these forms. According 
to Bromlei (1956), contrary to many other authors, the crossing of these species in 
nature was a very rare phenomenon in the past. No doubt this remains true even 
today. ^ 



136 

in general resemble sika deer more. Their fur in summer is spotted 
but the spots are yellow, not white, and less sharp. The antlers and 
tail resemble those of sika. The crossing of a male sika with a female 
Manchurian wapiti in a nursery invariably yields rust-colored off- 
spring without spots, similar to the female parent. Such forms are 
not encountered in nature (Menard, 1930). (V.H.) 

Taxonomy 

Within the genus Cervus the subgenus Ceruus, accepted here in a 
broad sense, represents the most specialized and progressive group. 
The evolutionary sequence of specialization proceeds as follows: 
from Axis to Rusa, Rucervus, and Przevalskium. Only in external 
characters are Russian sika similar to Axis deer (Indian spotted deer) 
with their permanent bright spotted coloration (in the past some 
researchers regarded and labeled the former Axis deer). Apart from 
several osteological characters, the former differ from the latter in 
the structure of the antlers; in Axis deer they invariably have three 
tines (intermediate forms absent). 

Within the subgenus sika represent a more primitive form than 
C. elaphus. This is borne out by the simpler structure of antlers 
(four tines, absence of bez tine and crown) and spotted coloration in 
adults. Yet the two species are undeniably quite close. Among its 
extremely diversified races С elaphus has forms without a bez tine, 
or without a crown, and also forms in which spotted coloration of 
the summer coat is not at all uncommon as an individual aberra- 
tion, or even usually present. Finally, some forms of this species are 
no larger than sika deer. The close affinity of these two species is 
underscored by their relatively frequent hybridization in nature. In 
captivity they freely interbreed in all combinations. (V.H.) 

Geographic Distribution 

These animals range over eastern Asia from the Ussuri region to 
southern China and northern Vietnam and some of the adjoining 
islands. 

107 Geographic Range in the Soviet Union 

The range in the Soviet Union comprises the extreme northeastern 
rim of the range of the species and occupies a very small territory in 
the extreme south of the Far East. 

The northern boundary of the range (reconstructed) is not fully 



137 




Fig. 34. Reconstructed geographic range of species of sika deer (Cervus 
nippon Temm.). V.G. Heptner. 

known. In the Ussuri valley and the western slopes of Sikhote-Alin 
mountains adjoining it the boundary extended almost up to the 
Bikin estuary (Maak, 1861)/^ i.e., roughly along 47°. 

The northernmost habitation of these deer on the eastern slope of 
Sikhote-Alin mountains is 46°00' (Abramov, 1928). Evidently these 
deer penetrated a few tens of kilometers in the north (Khutsin; 
Bromlei, 1956). The position of the boundary between these two 
points has not been established but there is no doubt that along 
Sikhote-Alin it curved strongly southward; deer were absent in the 
middle altitudes of the region. South of the above line deer were seen 
everywhere, including all the islands of Peter the Great Bay. The 
range extended beyond the boundaries of the Soviet Union in the 
south (into the Korean Peninsula) as well as to the west (into 
Manchuria). 

During the last century the range shrank notably and the north- 
ern boundary withdrew southward, at places very considerably. The 



'"These deer normally inhabit regions up to Iman; seen very rarely north of it, 
transgressions from the south v/ere recorded every year (Maak, 1861; Przewalski, 
1870). The extreme northern places of deer sightings were Bikhark mountains and the 
Diongo mountain chain (Maak, 1861) on the right bank of the Ussuri, slightly belovs^ 
present Nizhne-Mikhailov village on the LIssuri above the Bikin estuary. The nor- 
thernmost transgression occurred at Khat, slightly above Nora estuary on Ussuri 
(Maak, 1861). This location fell in the present-day Vidnoe village on Ussuri, below 
Bikinsk estuary and above Neolikha estuary (Nora), dropping into Ussuri on the left. 
In Bromlei's book (1956) the boundary according to data from Maak (1861) has 
been reconstructed incorrectly and cannot be properly deciphered. 



138 

108 main reasons for changes in the range were the extermination of 
animals by hunters and changes in topography. In recent years 
wolves have come to play an important role (see below). Currently 
the range (taking into consideration the occurrence of individual 
anirnals and small groups as well as transgressions) encompasses the 
far south of the Ussuri region, roughly up to 44° N lat. On the 
eastern slopes of Sikhote-Alin, in the region adjoining the sea, the 
range runs farther north. It extends as a narrow band northward up 
to Sikhote-Alin preserve and Khuntuez Bay (Abramov, 1954; G.F. 
Bromlei), i.e., roughly up to 45°20'-30'. The animals evidently 
reached even up to 46°. In this section no significant changes in the 
boundary occurred during the last century. 

Apart from a general withdrawal to the south, range reduction 
v^^as also evident in its division into several fairly isolated sections. 
The number of such sections (and also the number of animals in 
them) dropped rapidly and their isolation became complete. By 1950 
there were three such totally isolated sections on the mainland — 

109 Kedrovaya Pad' preserve (from the western bank of the Amur Gulf to 
west of Vladivostok), region of the former Sudzukhin preserve along 
the Sudzukh River to east of Suchansk, and Sikhote-Alin preserve. 
The number of animals in all these places is very small, not exceed- 
ing a few dozen (G.F. Bromlei). 

Deer are also present on islands in Peter the Great Bay — 
Askol'de, Putyatine, Rikord, and Rimsk-Korsakov (Abramov, 1954). 
These animals are descendants of wild deer but lead a semidomesti- 
cated life (for velvet antler recovery). Animals have not survived on 
the rest of the islands. 

Commencing from 1937 a small number of deer, taken from 
farms, were transferred to some preserves for purposes of acclimati- 
zation (Fig. 36). Deer were colonized in Il'men preserve (southwest of 
Chelyabinsk), Buzuluk (near Buzuluk), Kuibyshev preserve in Zhi- 
gulyakh, Mordov (southwest of Arzamas near Temnikov), Oka pre- 
serve (along the Pre River east of Ryazan'), Khoper (on the Khoper 
River near Borisogleb), and Teberdin preserve (in the northern Cau- 
casus on the Teberda River, tributary of the Kuban). There are no 
deer at present in the territory of Kuibyshev preserve but they con- 
tinue to survive in other places of acclimatization. Deer did not settle 
outside these sanctuaries or were present in extremely insignificant 
numbers. 



^ Deer detected in the upper courses of the Imansk around Sidatunsk village 
evidently represent escaped domesticated animals (G.F. Bromlei). 



139 




107 Fig. 35. Distribution of sika deer (Cervus nippon Temm.) in the Soviet Union. 
1 — reconstructed northern boundary circa 1850; 2 — boundary of range in last 
decade of nineteeth century and early present century; 3 — recent (1940's and 
1950's) habitats in Kedrovaya Pad', Sudzukhin, and Sikhote-Alin preserves. V.G. 

Heptner. 

Apart from the places listed above, deer have also been taken to 
Azerbaidzhan, Armenia, and several times to different places in Mos- 
cow district, around Vil'nyus, and some other areas. Such transfers 
were made by hunting organizations and the results are not yet 
definitive but so far hardly encouraging. Deer have likewise been 
imported into Borkuta in the floodplain of the lower course of the 
Dnieper and into several game farms of the Ukraine. These animals 
originated from Askaniya-Nova (for details, see below). 

Geographic Range outside the Soviet Union 

The range outside the Soviet Union covers the eastern part of north- 
eastern China — the mountain region adjoining the Ussuri region 
and Korea, part of the Sungari basin, Korea, and much of eastern 
and southeastern China. In the west the range includes southern 
parts of Jehol and western parts of Shansi, i.e., up to the meridianal 
course of the Hwang Ho, Henan, Hupeh, and Kwangsi. In the east 
the range extends up to the sea (specifically, it covers Shandong, 



'Chzhan-Guan-Tsailin, Khentei-Alin, and Van-Lun-Gou mountains (Baikov, 



1915). 



140 




108 Fig. 36. Some points of acclimatization of sika deer (Cervus nippon hortulo- 
rum) in the European part of the USSR, Urals, and the Caucasus (there are 
no deer at present in Kuibyshev preserve on Samarsk Luka). V.G. Heptner. 



Cheking, Fukien, and Kwangtung provinces). Farther away in the 
south the range extends into northern Vietnam (Tonking province, 
or Bakbo and Annam or Trunbo). 

Cheju-do (Qvelpart), Tsushima, Honshu, Hokkaido, Shikoku, 
Kyushyu, Yakushima, Ryukyu, and Taiwan Islands fall within the 
range. 

Over the greater part of China deer are totally extinct or 
extremely rare (in northeastern China and in the south they are still 
seen but few in number). 

Sika deer, particularly the Ussuri form, have been successfully 
acclimatized in New Zealand. (V.H.) 



141 

Geographic Variation 

The geographic variability of the species is slight within its rela- 
tively small range. A well-defined form inhabits the Soviet Union. 

Ussuri sika deer, С (С.) п. hortulorum Swinhoe, 1864 (syn. 
mantschuricus, tnantschuricus major, dybowskii, and hortulorum 
typicus), is the largest form of the species (for description, see above). 
Its range covers the Ussuri region. 

Outside the Soviet Union Ussuri sika deer are found in north- 
eastern China, Korea, and evidently the entire northern half of 
China (in the south up to the Hwang Ho). 

Three forms are distinguished outside the USSR: С п. kopschi 
Swinhoe, 1873 (from the Hwang Ho to the southern boundary of the 
range) is close to the Manchurian form and as large; С п. nippon 
Temm., 1838 (islands of Japan including probably Ryukyu) is prob- 
ably a distinct form characterized by very small size (height at 
shoulders about 95 cm); and C. n. taiouanus Blyth, 1860 (Taiwan). 
110 Some researchers distinguish up to nine forms of sika deer (in 
addition to those listed above: mantschuricus, mandarinus, grassia- 
nus, and pseudaxis from the mainland and keramae from parts of 
Ryukyu archipelago). Such distinctions are positively incorrect. At 
the same time a thorough analysis of geographic variability is pre- 
sently impossible since old descriptions are of little use, while new 
material is inaccessible since deer are extinct in much of the range of 
the species on the mainland. Theoretical concepts contravene recog- 
nizing a large number of races on the mainland. In any case Ussuri 
deer do not differ from Manchurian deer. 

Some earlier researchers (Przewalski, 1870; Maak, 1861) affirmed 
two species of sika deer in the Ussuri region. Such statements are 
based partly on an improper evaluation of individual variability, 
mainly in relation to the presence in the region of hybrids of this 
species and Manchurian wapiti (discussed above). That this is so is 
evident from the Chinese nomenclature cited by Maak (1861). (V.H.) 

Biology 

Population. Over the last 80 to 100 years the population of wild sika 
deer in the Soviet Union decreased several times, mainly as a result 
of their extermination by hunters in pre-Revolutionary Russia, 
especially during snowy winters, for example in the winters of 1877 
to 1878 and 1914 to 1915 (Yankovskii, 1882; Bromlei, 1956). At the 
end of the 1880's in the Far East the first antler farm based on sika 
deer was established. Such farms were initially filled with captured 



142 

animals and often poorly organized, leading to deer mortality. 
Finally, the gradual clearing of Primor'e forests, establishment of 
new villages, the appearance of a large number of wolves and hunt- 
ing dogs, and other factors also led to a reduction in the population 
of these wild ungulates. 

How greatly the population of sika deer decreased in the Far East 
can be judged from indirect indexes like changes in herd size. In the 
1860's and 1870's herds of 20 to 30 or even 40 to 50 animals were 
often sighted (Przewalski, 1870; Yankovskii, 1882). A group of 14 
deer has been observed only once throughout the existence of Sud- 
zukhin preserve (up to 1951), representing the main reserve of wild 
sika deer in the Soviet Union; all the remaining herds are no more 
than six or seven, and more often three or four strong (Bromlei, 
1956). 

Abramov (1930) estimated the number of wild sika deer at the end 
of the 1920's in the Far East at 900 to 1,100 (800 to 1,000 in the former 
Vladivostok region and up to 100 in Khabarovsk). In 1949 they did not 
exceed 300 (Bromlei, 1956), of which 160 were in the territory of 
Sudzukhin preserve, 20 in Sikhote-Alin, 30 each in Suputinsk and 
Kedrovaya Pad', and 60 outside the preserves. After Sudzukhin pre- 
serve was closed down in 1951 the number of wild sika deer 
decreased further. 

In 1938 a total of 240 sika deer were transferred from Far East 
reindeer farms to Il'men preserve (Trans-Urals) and other sanctuar- 
ies in the European part of the USSR and set free. They have flour- 
ished and at places penetrated into regions adjoining the preserves 
(not more than several tens of kilometers from the points of release) 
but their number has only doubled as of now. In most of the new 
regions the snow cover has proved too severe for their survival. In 
1953 to 1954 in these preserves and adjacent areas sika deer num- 
bered as follows: Teberdin— 21 to 36, Khoper— 206 to 209,^' Oka— 70 
111 to 99, Mordov — 151, Il'men — 20 to 35, and Kuibyshev preserve and 
Buzuluk Bor a few solitary animals, i.e., a total of 500 deer. Maxi- 
mum population growth has been seen in Khoper (7 to 10 times), 
Oka (3.5 times), and Mordov (2.5 times) preserves. In the other pre- 
serves the number of deer did not increase or, in fact, decreased 
(Il'ina, 1956). In Teberdin preserve deer had almost totally disap- 
peared by 1956; at present, they are found only outside the preserve, 
in the much lower parts of Karachai autonomous region (Arens, 1957; 

^°In 1957 the preserve was reopened. 

^'Up to 300 deer were counted in 1955 in Khoper preserve and neighboring 
forests (D'yakov and Aleinikov, 1956). This figure appears somewhat inflated. 



143 



Inyakova, 1957). In Kuibyshev preserve sika deer have disappeared 
altogether (I.V. Zharkov). 

Small numbers of sika deer were brought into different regions of 
Moscow^ district between 1933 and 1955: 6 animals were released in 
1952 in Khuzinsk region of Azerbaidzhan Soviet Socialist Republic, 
and 20 in 1953 in Vedinsk region of Armenian Soviet Socialist 
Republic (Il'ina, 1956). As early as 1909 sika deer were imported into 
Askaniya-Nova where they are present even today. In 1941 about 80 
deer were set free in the Dnieper division of Askaniya-Nova preserve 
at Borkuta; deer in the Dnieper floodplain (on the left bank) have 
multiplied and their number now is appreciably greater (Salganskii, 
1952). In 1956 over 100 deer were imported from Askaniya-Nova into 
game farms in the Ukraine. Deer have also been set free in the region 
of Black Sea preserve. Recently sika deer (24 animals from Altai state 
farm) were released near Kaunas (Lithuanian Soviet Socialist 
Republic) and in 1955 produced offspring (Ivanauskas, 1957). 

For the Soviet Union as a whole the population of wild sika deer 
is probably 1,000 or more. 






Fig. 37. Sika deer in summer coat. Il'men preserve. 
Photograph by N.N. Nemnonov. 



144 

Habitat. In the Primor'e region sika deer inhabit mainly broad- 
leaved oak forests of the Manchurian type and rich undergrowths, are 
rarely seen in mixed nut pine- broad- leaved forests (usually not 
above 500 m), and do not penetrate nut pine-conifer taiga; they 
avidly feed in growing burned-over forests. Sika deer find best condi- 
tions at places where the total winter precipitation in the form of 
snow does not exceed 800 to 1,000 mm, the first snowfall commences 
relatively late, the total number of days with snow cover averages 
less than 45, and the height of the snow usually does not exceed 25 to 
30 cm. Of particular interest to deer are the southern and southeast- 
ern slopes of mountain ranges along the sea, where the snow cover 
usually does not last longer than five to eight consecutive days since 
it is soon washed away by rains. Preferred habitats are those with 
rugged topography and abundant streams. Females and young deer 
are found in large numbers slightly lower on slopes and closer to the 
sea compared to adult males (Abramov, 1929 and 1939; Bromlei, 
1959). 

Sika deer released in Trans-Urals and the European part of the 
USSR find conditions there incomparably more difficult in winter, 
112 as the snow cover persists for up to 150 days (Il'men preserve), its 
height often exceeds 60 cm (Il'men and Mordov preserves, Zhiguli, 
and Buzuluk Bor), and minimum temperature at some places 
(Buzuluk Bor) reaches — 49.5°C (Arsen'ev, 1949). Deer in these 
regions are confined mainly to deciduous and mixed forests; in win- 
ter in Il'men and Oka preserves, when the snow is deep, they are 
usually confined to pine groves (Il'ina, 1956). In Teberdin preserve • 
sika deer sometimes penetrate to a height of 1,600 to 1,700 m, which 
is not characteristic of deer in the Primor'e region. 

Food. In the Primor'e region wild sika consume no less than 129 
plant species. Trees and shrubs play the most important role, espe- 
cially in summer; their proportion in the diet of deer reaches 70% (by 
volume) (Bromlei, 1956). Among the main food items are acorns, 
leaves, buds and slender branches, oak shoots, Manchurian aralia, 
and lime. Lespedeza, acanthopanax, Amur zovk, Manchurian nuts, 
and Amur grapes are also preferred items. In spring elm, maple, ash, 
and sedge, and in summer umbelliferous and other plants become 
dietary items. Deer gnaw tree barks other than oak (which is totally 
inedible) mainly in the second half of winter when the snow is 
heavy. Under very similar conditions branches of willow, bird 
cherry, and alder also serve as food. Deer feed on acorns in winter 

"Manchurian wapiti consume thicker branches as they have a thickened skin on 
the Hps; sika deer have no such thickened skin (Bromlei, 1956). 



145 

and continue to dig them up from under snow up to 30 cm deep and 
when the acorns are abundant, even when the snow is 40 to 50 cm 
thick. They feed well not only on green but also on dry oak leaves 
clinging to branches as well as those which have dropped if they are 
not rotten. Deer dig up leaves from under the snow. On the sea coast 
they look for eel grass and laminaria, attracted by their saltiness; 
deer usually consume them only in winter and simply chew them in 
summer. Unlike other ungulates these deer usually feed poorly on 
grassy substances and arboreal lichens^^ (Vendland, 1938; Bromlei, 
1956). 

In preserves in the European part of the USSR and in Trans- 
Urals sika deer consume over 390 varieties of plants, of which only 
about one-eighth are the same as those eaten in the Primor'e region; 
15 species are consumed avidly and 173 well. Some species of plants 
consumed well in preserves in the European part of the USSR and 
Trans-Urals are poorly consumed in the Primor'e region and vice 
versa. Compared to the Primor'e region, the role of forest shrubs in 
the summer diet is significantly less; deer at this time of year feed 
mainly on tall grasses (Il'ina, 1956). 

In Il'men preserve the consumption of 143 plant species has been 
recorded (Averin and Ushkov, 1947), in Mordov — 36 plant species 
(K.N. Nikitin), Khoper— 132 (D'yakov and Aleinikov, 1956), 
Kuibyshev — 154 (A.N. Goncharova; and others), farms of Buzuluk 
Bor — 80 (Darkshevich, 1939), and around Moscow — 63 (Matyushin, 
1954). In the Kuibyshev preserve steppe regions where deer have 
settled (Zhiguli) they consume in early spring 110, in winter 60, in 
summer 55, and in autumn 34 plant species. Warty barked spindle 
tree [Euonynus verrucosus^, maple, rowan, oak, pine shoots, etc. are 
consumed more avidly than others (A.N. Goncharova). In Teberdin 
preserve acorns and beechnuts are well consumed (when available, 
deer dig them up from under snow 50 to 60 cm thick) and branches 
of hazel, oak, aspen, willow, elm, and especially spindle tree. Those 
of special interest from among herbaceous plants are cow parsnip, 
spirea, lady's mantle, cinquefoil, etc. (KurapovaandStepanov, 1940; 
Knyazev, 1946). In Oka preserve in winter deer feed mainly on the 
branches of willow, brittle buckthorn, oak, and aspen (Il'ina, 1956). 
In Il'men and Mordov preserves, when the snow is 30 cm thick, deer 
almost totally consume hay from places of cattle grazing and stocks 
dropped accidentally on the roadside. When food is inadequate 
113 hungry deer in Mordov preserve consume large quantities of young 

In farms around Moscow and in Khoper preserve wild deer take well to arboreal 
lichens (Matyushin, 1954; D'yakov and Aleinikov, 1956). 



147 



pine shoots and gnaw at pine and spruce bark (P.M. Reshetnikov; 
F.D. Shaposhnikov; and others). 

In Suifunsk deer farm domesticated sika deer consume 179 plant 
species from 37 families (of the total number of 364 species of forest 
plants in southern Primor'e), including trees, shrubs, herbage, 
legumes, grasses, aspen, horsetail, and ferns. Oak, lime, and briar 
{Smilax) are more avidly consumed, 52 species consumed well, and 
72 satisfactorily. Among the plants consumed are aromatics with a 
pungent smell and those which are poisonous to cattle (Ryabova 
and Saverkin, 1937). Thus the stenophagy and poor adaptability of 
sika deer in food selection reported by some researchers is not 
confirmed. 

A sika male weighing 100 kg requires 2.0 kg food containing 0.20 
to 0.25 kg digestible proteins per day, while aduh females of different 
ages require 1.6 to 2.4 kg food containing 0.14 to 0.22 kg digestible 
proteins (Mirolyubov and Ryashchenko, 1948). 

Deer visit some warm mineral springs and sometimes drink sea 
water. In search of mineral matter they lick algae and other marine 
materials washed ashore (holothurians, pebbles, etc.) and also ashes 
(Bromlei, 1956). Deer visit artificial salt licks only in some regions 












Fig. 38. Herd of sika deer in Maikha state farm. Primor'e region, 1953. 
Photograph by N.N. Nemnonov. 



148 

where they have been acchmatized. 

Home range. Some lone animals and herds of sika deer have their 
own home range, the size of which for lone animals rarely exceeds 
100 to 200 hectares. In the period of rut a male with four or five 
females is confined to an area of about 400 hectares, and a much 
larger herd (14 to 16 animals) to a region of 800 to 900 hectares 
(Bromlei, 1956). In winter in Primor'e region, especially when the 
snow cover is very thick, many deer survive for weeks in an area 
comprising several tens of hectares or even smaller, for example, 
wandering along unfrozen streams and feeding on trees along the 
banks. In winter the daily wandering of deer in Sudzukhin preserve 
often exceeds 300 m (G.F. Bromlei); in Mordov preserve when the 
snow cover is about 50 cm deep wandering averages 1 ,500 m for a male, 
114 but only 500 m when the animal feeds on hay (F.D. Shaposhnikov). 
In Oka preserve, when the snow is not more than 30 cm thick, daily 
travel does not exceed 2.0 to 2.5 km; when the snow is very thick or 
has a frozen crust daily travel is limited to 0.6 to 1.5 km (Il'ina, 1956). 
In the total area of 339,000 hectares of Sudzukhin preserve, sika 
have been sighted in summer only in 42,000 hectares and in winter 
in 15,000 hectares. In summer, depending on the total population of 
sika deer in the preserve, their density has varied in different years 
from 3.5 to 12 animals per 1,000 hectares; density is higher in winter. 
This value under conditions of Primor'e region is closer to maxi- 
mum for the species (Bromlei, 1956). 



W^''!^''SVM1 




Fig. 39. Track of sika deer in Mordov preserve. March, 1954. Photograph by 

G.I. Il'ina. 



149 

In the Far East, when farm deer are fed hay and twigs in winter 
and spring, about 1.0 to 1.5 hectares of pasture is adequate per 
animal, but in this process the vegetation of the area allotted is 
gradually depleted.^'' If the vegetation is to be maintained in good 
condition, each deer should be allotted 10 to 15 hectares of pasture. 
Such an allotment is uneconomical, however, as this ratio would 
greatly enlarge the farm area required (Mirolyubov and Ryash- 
chenko, 1948). 

Daily activity and behavior. In summer in Primor'e region sika 
deer feed mainly in the morning and evening; the winter rhythm of 
feeding is not so well regulated. The mobility of deer is greatly 
hindered during snowfall and the animals stay bedded down; on 
clear days, during frosts, their activity notably increases. In winter, 
when winds are gusty, the animals look for shelters in sections of 
dense forest and bed down in becalmed places. In preserves in the 
European part of the USSR the daily rhythm of sika deer has yet to 
be fully ascertained. 

The weight load carried by sika deer per sq cm area of hooves is 
830 to 1,140 g, or an average of 960 g (Bromlei, 1956), which is 
significantly greater than the corresponding values for other ungu- 
lates of the Soviet Union. The heavy load on the hoof of the animal 
and its relatively small structure makes it very difficult for it to reach 
food and move about on thick snow. At a snow cover of 60 to 65 cm 
thickness deer progress only by hops and soon exhaust themselves. 
In the absence of snow they are capable of running swiftly for quite 
some time and easily clear hurdles up to 1.7 m high (Bromlei, 1956). 

Deer swim well. Instances are known of deer swimming 10 to 12 
km in the sea, for example from Putyatinsk to AskoI'd Islands 
(Abramov, 1954). In Oka preserve sika deer stand in water to protect 
themselves from blood-sucking flies (Il'ina, 1956). 
115 The organs of hearing, vision, and smell are well developed in 
sika deer. 

After rut adult males isolate themselves and live in winter in 
small groups. Herds of females include young deer of both sexes not 
older than two years. The gregarious tendency is generally more 
prominent in winter, especially in years of abundant acorn crops 
(Abramov, 1954). In Oka and Khoper preserves herds of up to 23 to 
25 animals have been recorded in winter (Il'ina, 1955). 

^''When deer density is even higher, vegetation is rapidly exhausted. Under condi- 
tions of unsystematic grazing and a deer density of one animal per 0.5 hectare, up to 
80% twigs and 25 to 50% herbaceous vegetation of the park were depleted in one year's 
time (Ryabova and Saverkin, 1937). 



150 

Seasonal migrations and transgressions. Nearly the entire popu- 
lation of sika deer in Primor'e region winters on the eastern slopes of 
Sikhote-Alin. Commencing from December deer congregate on the 
southern and southeastern parts of the slopes and most move closer 
to the sea by descending into the broad-leaved forest belt. Most of the 
animals are confined to open valley regions, well-protected from 
cold westerly mountain winds blowing down the slopes (Bromlei, 
1956). At the end of the 1940's on the large western slope of Sikhote- 
Alin there was only one small wintering site; in Sikhote-Alin sanc- 
tuary animals live along the steep parts southward (V.D. Shamykin). 

In Primor'e region where heavy snowfalls are not rare in Febru- 
ary and March, deer are compelled to migrate even closer to the sea. 
In this period most remain within 3 km of the sea; the population is 
scattered into several isolated groups living on relatively steep and 
less snowy (up to 30 cm) slopes. When the snowfall is particularly 
heavy, deer find refuge alongside tiny unfrozen streams and in lit- 
toral zones (Bromlei, 1956; and others). In winters of moderate inten- 
sity, even a minor snowfall can cause the movement of deer into less 
snowy sections of the region. After a few days the animals again 
scatter widely (G.F. Bromlei). In Sikhote-Alin preserve sika deer 
represent a more settled species compared to elk [moose] and Man- 
churian wapiti (V.D. Shamykin). 

In May deer move into summer habitats, often selecting forest 
sections with dense undercover. When blood-sucking flies and ticks 
are numerous, some deer remain close to the sea coast and emerge 
into open, windy regions (Bromlei, 1956; and others). 

In the European part of the USSR sika deer are characterized by 
seasonal migrations. ^^ Such migrations are best seen in Khoper pre- 
serve; during the spring floods much of the Khoper becomes water- 
logged and some deer move into the adjacent Borisogleb forest zone. 

Reproduction. In Sudzukhin preserve rut commences between 
September 29 and October 11, but in 1944 a bellowing bull was 
heard on September 10;^^ the period of mass and most intense bel- 
lowing extends from October 7 to 23, lasting in various years for 
four to seven days. Rut ceases by November 5 to 8, more rarely at the 
end of October. The period of rut on the whole continues for 25 to 35 
days but each adult female responds for less than a week. During the 
years when acorns are available rutting is very energetic. In October 

"ll'ina (1956), without proof, contests this fact. 

"in his conclusions Bromlei (1956) erroneously states (p. 213) that rut commen- 
ces September 10 even though he has given correct dates earlier in the text (pp. 183 
and 184). 



151 

adult males move with small (three to four animals) harems of 
females. Compared to Manchurian wapiti, rut in sika deer proceeds 
more gently and no combats have been observed between males." In 
the morning and evening adult males emit a whistling sound which 
ends in a hoarse howl. Usually the howl issues from those areas 
where deer have stamped out small (4 m"^) mating sites with their 
hooves. A single male usually makes six or seven such sites. In the 
period of rut males eat little and lose 20 to 25% of their normal 
weight (Bromlei, 1956). 

Deer, at least females, may become sexually mature from 1.5 
years, but most females do so in their third, rarely second year. Males 
116 enter rut not before the third or fourth autumn of life; males 1.5 
years of age do not bellow (Bromlei, 1956; and others). 

Among acclimatized deer the period of intensive rut roughly 
coincides with that in Primor'e region, but on the whole is more 
prolonged. In Il'men preserve rut occurs from the end of September 
to the end of October (climax in mid-October); some individual 
males bellow even in early December. Usually rut is interrupted by 
snowfall and cold (P.M. Reshetnikov). In Khoper preserve, based on 
observations covering a period of 18 years, rut commences from 
September 18 to October 20, the very last bellowing males being 
heard on December 15. In an overwhelming majority of deer rut 
occurs in October and extends for an average of 34 days (D'yakov 
and Aleinikov, 1956). In some preserves bellowing males are heard 
even in January. In Moscow region most captive deer enter rut from 
mid-October to mid-November (Matyushin, 1954) and in Teberdin 
preserve from October 5 to 10 through mid-November (Kurapova 
and Stepanov, 1940). 

The normal duration of gestation of sika deer is around 7.5 
months; often it is a few days less (Mirolyubov and Ryashchenko, 
1948; Matyushin, 1954). In Primor'e region barren females are 
encountered very rarely among wild deer. In Sudzukhin preserve the 
earliest case of normal fawning is seen between April 20 and 30 and 
the last on June 18; most fawning occurs mid-May. A female with 
more than one fawn has never been encountered but once two 
embryos were found in a dead female on dissection (Bromlei, 1956). 

In most acclimatized deer fawning occurs in June; some individ- 
ual females may fawn in July-August or even in September- 

"in preserves in the European part of the USSR where the sex ratio is evidently 
different, intense combats are common between males; in Khoper preserve instances 
of male deaths in the course of combat have been recorded (D'yakov and Aleinikov, 
1956). Intense combats between males are also common in farms where deer are raised 
for their velvet antlers. 



152 

November. Three instances of twins, all in Il'men preserve, were 
observed over a period of 16 years, starting from the commencement 
of acclimatization (Arsen'ev, 1949; Il'ina, 1956). 

In velvet antler husbandry rut and fawning of sika deer occur at a 
much later period than in wild deer in the Far East and, generally, 
less evenly; some females become sexually active only in early win- 
ter. A single male mating with 5 females is considered normal; 
strong bulls can mate with 10 to 20 females. Barrenness is considera- 
bly more than in wild forms and, in some farms, as high as 30 or 
even 70%. Usually, there is a single fawn; twins are rare. Some 
females, commencing from the age of 1.5 years, fawn annually. 

Growth, development, and molt. Newborn deer remain helpless 
for the first few days, lying down most of the time; mothers graze in 
the vicinity. Most fawns suckle for four to five months. In Sudzukhin 
preserve a lactating female was observed in November and one was 
once caught in January. Ten to 20 days after birth the young com- 
mence grazing independently. They remain with the mother until 
the spring of the following year, however, and sometimes even 
longer (Bromlei, 1956; and others). 

Newborn bucks of captive animals weigh 4.75 to 7.32 kg (12 
observations) and does 4.2 to 6.2 kg (9 observations). By age 9 to 12 
months wild male fawns weigh 49 to 59 kg and female fawns 48 to 55 
kg; at the end of 16 months weight is 70 to 75 and 65 to 70 kg 
respectively; in winter fawns gain almost no weight. After two years 
weight increase slows down. Around that age males start outstrip- 
ping females noticeably in growth. The maximum weight of males 
is recorded at 7 to 10 years and of females at 4 to 6 years. In Sudzuk- 
hin preserve the maximum, correctly established, weight of a male 
was 131 kg and that of a female 85.7 kg; corresponding weights in 
velvet antler farms were 148 and 112 kg respectively (Mirolyubov and 
Ryashchenko, 1948; Matyushin, 1954; Bromlei, 1956). 

At age 1.0 to 1.5 years milk incisors are replaced by permanent 
ones and at age 2.0 milk molars also replaced. By the tenth month 
small "tuberosities" (3.5 cm) are visible in males at the place of 
future antlers; by April the first tiny unbranched antlers have begun 
to grow. "Shilniks" bear such antlers around the first year of life (up 
to May-June of the following year). As soon as these are shed 
branched antlers, or the first velvet antlers, begin to grow. In 
nurseries the maximum growth of antlers is achieved at the age of 10 
117 to 12 years. In Primor'e region adult deer shed antlers at the end of 
April or in May; at the end of May-June young velvet antlers begin 
to appear. In September deer rub the velvet off their antlers (Brom- 
lei, 1956). With a month's delay, or more rarely around the same 



153 

time as in Primor'e, the shedding of old and the growth of new 
antlers take place in deer in preserves of the European part of the 
USSR, in the Caucasus, and in Trans-Urals (Kurapova and Ste- 
panov, 1940; Il'ina, 1956). 

The juvenile spotted coat is discarded after the autumn molt. In 
the Primor'e region spring molt of adults commences in the first 10 
days of March (hair of the beard falls first), peaks in the latter half of 
April to early May, and ceases in the last 10 days of May. In the sick 
and ailing molt is delayed. The summer coat is sparse, shorter than 
the winter one, and almost devoid of underfur. Winter hair begins to 
grow perceptibly at the end of August and by mid-September, or 
slightly earlier, deer acquire their winter coat and hair growth is 
negligible thereafter (Bromlei, 1956; and others). In preserves of the 
European part of the USSR spring molt of deer occurs after a slight 
delay compared with the Far East; moreover, it is more extended in 
the above preserve (Il'ina, 1956). 

According to observations made in velvet antler farms the maxi- 
mum life span of sika deer is 18 to 21 years; females older than 15 
years can still breed. In nature deer older than 11 to 14 years usually 
do not survive as they are killed by predators or die from other causes 
(Mirolyubov and Ryashchenko, 1948; Bromlei, 1956; and others). 

Enemies, diseases, parasites, mortality, competitors, and popula- 
tion dynamics. Among various factors causing death of sika deer, 
predators, especially gray wolf, are of maximum importance. In 
Mordov preserve from 1948 to 1952 the death of 111 sika was 
recorded, of which 50 (45%) were caused mainly by wolves and partly 
by lynxes, 25 were due to emaciation and disease, 4 to traumatic 
injuries, etc. (F.D. Shaposhnikov). In Oka preserve 27 instances of 
deer mortality were reported from 1938 to 1951, of which 12 (44%) 
were caused by wolves, 6 by dogs, 1 by lynxes, 3 by disease, and 4 by 
drowning in floodwaters flowing under ice (Kozlov, 1954). In 
Khoper preserve from 1939 to 1947 no less than 50 deer were des- 
troyed by wolves, of which 11 were killed in 1943 alone (Arsen'ev, 
1949; and others). During the 17 years of acclimatization in preserves 
of the European part of the USSR, the Caucasus, and Trans-Urals, 
383 instances of death of sika deer have been registered; of these, 
45.7% were caused by wolves, 0.5% by lynxes, and 3.1% by stray dogs 
(Il'ina, 1956). 

In Sudzukhin preserve 5 to 30% of the population of sika deer are 
killed annually by wolves (gray). In the feces of wolves (gray; 59 
samples) the remains of sika deer were detected in 16.9% in January 
and February, 39% in March, 22.2% in April, and in much smaller 
numbers at other times. Thus wolves cause maximum damage to 



154 



sika deer of Sudzukhin preserve in the second half of winter when 
the snow cover is usually maximum and frozen snow crust forms 
often. This is also the time of massive ice formation on rivers and 
deer slip on ice helplessly (Bromlei, 1956). In attempting to save 
themselves from wolves deer of Sudzukhin preserve often run into 
dense shrubs, penetrate relatively deeply into rivers, or find refuge in 
the sea, where they remain sometimes for 30 to 40 minutes and swim 
400 to 1,000 m or even farther away from the coast. On frosty, slushy 
days many lose their stamina, their lungs become inflamed, and 
death ensues; pregnant females often abort (Bromlei, 1956). In Ked- 
rovaya Pad' preserve 83 deaths of sika deer were recorded from 1926 
to 1936; deaths were due mainly to red and partly to gray wolves 
(Mirolyubov and Ryashchenko, 1948). In Khoper preserve several 
deer have been killed by wolves in the least snowy winters, when the 
lakes are not fully covered with ice and deer slip and often fall into 
them (D'yakov and Aleinikov, 1956). 

Apart from the predators mentioned above, sika deer are also 
threatened by Far East leopards and tigers, but their numbers are 
insignificant. Fawns, especially up the age of two months, are threat- 
ened by bear. Far East leopard cat, red, fox, and yellow-throated 
marten. Instances of attack by predatory birds are not known. Sud- 
118 zukhin preserve has registered six deaths of deer caused by stray dogs 
(Bromlei, 1956; and others). 

Sika are susceptible to foot-and-mouth disease, rabies, carbuncu- 
lar emphysema, necrobacillosis, hemorrhagic septicemia (pasteurel- 




Fig. 40. Male sika deer dead on slippery river ice. Khoper preserve. October 
1954. Photograph by G.I. Il'ina. 



155 

losis), Siberian uclers, tuberculosis, leptospirosis, actinomycosis, 
ringworm, and coccidiosis. From 1932 to 1940 in the velvet antler 
farms of Primor'e region 470 deer died of necrobacillosis and 134 
from hemorrhagic septicemia (Mirolyubov and Ryashchenko, 1948; 
Vysotskii and Red'kina, 1954; and others). Instances of deer mortal- 
ity due to hemorrhagic septicemia have also been reported from Oka 
and Teberdin preserves and as a result of foot-and-mouth disease 
from Buzuluk Bor (Il'ina, 1956). 

In Primor'e region and in places of acclimatization only a few 
species of helminths have been detected in wild sika. In Sudzukhin 
preserve 75% of adult deer were once infected with the liver fluke 
Dicrocelium lanceolatum (Bromlei, 1956). Among animals held in 
feedlots 4 species of trematodes, 3 species of cestodes, and 29 species 
of nematodes have been recorded (Abramov, 1954). Between 200 and 
500 ticks {Dermacentor, Ixodes, and others) were found in early June 
in some wild sika deer. Deer also suffer from gnats (mainly Simulum 
maculatum), biting midges, horseflies, sucking lice (Lipoptena 
cervi), body lice, and other ectoparasites. Severe parasitism by naso- 
pharyngeal gadfly has been established in deer in feedlots but wild 
deer are not invaded by it (Bromlei, 1956). 

In snowy and severe winters in Primor'e region (1877-1878, 
1914-1915, 1928-1929, 1934-1935, 1938-1939, 1941-1942, and 1947- 
1948) a considerable number of deaths of deer due to emaciation, 
predators, and poachers were registered. The winter of 1947-1948 
was particularly damaging when, in Sudzukhin preserve, up to 50% 
of all the deer died; in some adjacent regions mortality was 100%. 
When the snow cover (over 40 cm on relatively steep slopes) does not 
remain longer than a month, deer can withstand such winters relatively 
well (Bromlei, 1956). 

Roe deer and Manchurian wapiti occupy the same habitats as 
sika; however, in view of the relatively low population of ungulates 
in the regions of their coexistence in the Far East, competition 
between these species is negligible. In Il'men preserve, of the 125 
plant species which are satisfactorily and well consumed by sika 
deer, roe deer consume about 90 (Averin and Ushkov, 1947). 

Captive sika deer can interbreed with other deer (maral, Manchu- 
rian wapiti, and Crimean red deer); in nature, such instances are 
extremely rare. 

The population pattern of wild sika deer in Primor'e region can 
be judged from the following data: of the 38 deer which died in 
Sudzukhin preserve, 55% were not older than two years and only 5% 
older than 10 years (Bromlei, 1956). Two instances are known when 
15 sika protected from wolves grew to a herd of 2,000 to 2,500 ani- 



156 

mals in 30 years (Sidima Peninsula and Askol'd Islands). In Oka pre- 
serve the annual herd growth averaged 20% (Kozlov, 1954) and in 
Khoper 14.6 and 31.1% from 1938 to 1953 and 1954 respectively 
(D'yakov and Aleinikov, 1956). In Askaniya-Nova 80 to 93% of new- 
borns survive up to six months (Salganskii, 1952). 

Among embryos and newborn animals males occur about 1.5 
times more often than females. After a few months this ratio changes 
in favor of females because of high male mortality. In 1945 in Sud- 
zukhin preserve 216 females and 54 males of sika deer were counted; 
thus, the sex ratio was 4:1 in favor of females (Bromlei, 1956). In 
Oka preserve the ratio of male to female is 1:3 (Il'ina, 1956). 

Field characteristics. In growth sika deer lag noticeably behind 
red deer Curvus elaphus, especially Manchurian wapiti. Only some 
individual large male sika deer attain a height of 120 cm at the 
withers, while even female Manchurian wapiti reach 118 to 135 cm, 
on the average 128 cm, at the withers. The summer coat is character- 
ized by distinct spots. In size of hoofprints and feces sika deer fall 
between Manchurian wapiti and roe deer (Formozov, 1952). The 
pace is 45 to 65 cm, which on trotting lengthens up to 75 to 110 cm, 
when jumping up to 3 to 3.6 m, and when jumping over ravines up 
to 6 m. Sika strip the bark of trees in narrow longitudinal bands and 
not continuously as done by Manchurian wapiti; they do not remain 
for long near a single tree (Bromlei, 1956). Before lying down sika 
rake up the snow (if it is wet) or lie directly on it (when the snow is 
high and dry). Stamped tracks are characteristic of sika deer. 

Females emit a loud whistling call which terminates in a hoarse 
rattle. Young males also whistle in a similar manner, but without 
the rattle at the end. Adult males usually call only during rut (Brom- 
lei, 1956). 

The summer fur of adult males and females is similar; the main 
shade is rust with white spots (in females the color is somewhat 
lighter). In winter differences are more distinct: in females the gen- 
eral background is light gray, and in males, olive-brown and gener- 
ally even darker. (A.N.) 

Economic Importance 

120 In view of the low population of wild sika deer their hunting has 
long been prohibited. In the nineteenth century most deer were 
caught on rocky littoral shoals or islets in long enclosures in which 
pits were dug. Russian game hunters used to chase the deer toward 
the sea and then shoot them. 

The meat of this deer is good; the quality of male meat is at its 



157 




Fig. 41. Hoofprint of male sika deer (magnified). Primor'e region. June, 1928 
(from Formozov, 1952). 

best in September, i.e., before rut, and the female meat in September 
to November. Hides can be used to make chamois, rugs, and various 
articles. Young unossified velvet antlers^^ are considered the most 
valuable deer product; from these antlers is prepared the powerful 
tonic, pantocrine. The curative properties of velvet antlers of sika 
are rated superior to those of other deer and hybrid forms. Studies 
conducted on a small amount of material in Mordov preserve 
revealed that the velvet antlers of deer acclimatized there possessed 
only little "activity" (Arsen'ev, 1949). This conclusion is of utmost 
importance; without testing, it cannot be applied to other regions of 
deer acclimatization. The largest and best velvet antlers are found on 
deer 7 to 8 and 11 and 12 years of age. 

The great demand for velvet antlers and their high value in the 
Chinese market promoted development in the Soviet Union of a 
special branch of animal husbandry — raising sika deer to obtain 
velvet antlers. In the Far East the confinement of deer in paddocks 
was instituted at the end of the 1880's. Such small individual farms 



^*Velvet antlers of stage V, before the commencement of ossification of the carti- 
laginous mass, are valued most. In wild deer of Sudzukhin preserve this stage corre- 
sponds to the period June 15 to 20 (Bromlei, 1956). 



158 





Fig, 42. Winter feces of female (top) and male (bottom) sika deer (natural size). 
Sudzukhin preserve. Sketch by G.F. Biomlei. 



have greatly multiplied in present-day Khasansk, Suchansk, and 
Shkotov regions, and in southern Ol'ginsk and Terneisk regions. 
121 Large parks also existed, for example, Yankov park on the Sidima 
Peninsula (Abramov, 1954). In 1928 deer farms and parks were 
nationalized and formed the basis for large state farms in which deer 
were raised to obtain velvet antlers. In the mid 1930's over 8,500 sika 
deer were held in Primor'e state farms and 1,000 in collective farms 
(Ryabova and Saverkin, 1937). Sika were imported into nurseries of 
the Irkutsk region and the Altai (initially in 1933 into Shebalino, 
and later into Kaitanak, Nizhnii Uimonsk, and the upper courses of 
the Katun) and held in common paddocks with maral or separately. 
In the mid-1 950's there were up to 20,000 sika in such nurseries 
(Il'ina, 1956). 

As a result of diminished demand for velvet antlers in the inter- 
national market future prospects for velvet antler farming are not 
clear. Neither are the prospects clear for such products as three- to 
four-month-old embryos removed from the womb of female deer, 
tails, penises, etc., which have a market in China. The importance of 



159 

other marketable products of sika (sinews, ossified horns, and leg 
bones) is relatively insignificant. 

In the past, when these deer were far more numerous, they threat- 
ened plantations in some remote forest segments. At present, how- 
ever, they must be strictly conserved everywhere. It is also extremely 
desirable to reacclimatize sika deer in regions of Primor'e where they 
existed in the past but were subsequently exterminated. It is also 
possible to release these deer on some islands near Vladivostok. 
(A.N.) 

4. TRUE, WELLBORN [RED] DEER 
Cervus (Cervus) elaphus Linnaeus 1758^^ 

1758. Cervus elaphus. Linnaeus. Systema Naturae, 10th ed., vol. 1, p. 

67. Southern Sweden. 
1717. Cervus elaphus hippelaphus. Erxleben. Syst. Regni Anim., 

vol. 1, Mammalia, p. 304. Ardennes. 
1777. Cervus elaphus canadensis. Erxleben. Syst. Regni Anim., vol 1, 

Mammalia, p. 305. Eastern Canada. 
\MQ. Cervus maral. Ogilby. Rep. Council Zool., p. 22. Nomen 

nudum. 
\^^\. Cervus affinis. Hodgson. Journ. Asiat. Soc. Beng., vol. 10, pt. 

2, p. 721. Chumbi valley in Sikkim. 
IS50. Cervus maral. Gray. Knoiasly Menagerie, pis. 38 and 39. Cas- 
pian province of Iran. 
1867. Cervus xanthopygos. Milne-Edwards. Ann. Sc. Nat. Zool., vol. 

8, p. 376. Peking environs (possibly northern Manchuria, 

Ussuri). 



Usually the European form called the red deer is considered a separate species, 
distinct from izyubr, maral, American wapiti, Bukhara deer, and other forms, which 
are also often treated as separate species. After it was demonstrated that all these forms 
shared a common species background (Heptner, 1940; Heptner and Tsalkin, 1947), and 
this viewpoint became generally acknowledged, it became more advantageous to 
combine these species under the Russian name "true deer," which is distinct from 
spotted and other deer. The popular names "maral," "Manchurian wapiti," etc. are, 
of course, best retained for some individual varieties or groups of varieties. It is 
meaningless to call the Russian maral by the American name wapiti, as is sometimes 
done. 

Both these names are artificial and bookish. The Russian popular name of this 
species in the western part of the geographic range, in the Caucasus, and in Central 
Asia is "deer," in Siberia "maral," and in Trans-Baikal and the Far East "izyubr". 
Among hunters the adult male is called a bull, the adult female "olenukha" or 
"maralukha," and a two-year-old deer with simple antlers "saek" or "spichak". 



160 

1872.(1873).^° Cervus maral var. songarica. Severtsov. Vertical and 

Horizontal Distribution of Turkestan Animals, p. 109. Eastern 

part of Trans-Iliisk Alatau (upper course of Turgena).^' 
122 1872.(1873). Cervus maral var. sibirica. Severtsov. Vertical and 

Horizontal Distribution of Turkestan Animals, p. 109. Type 

locality has not been directly indicated but it is clear from the 

text that the Altai should be regarded as such. The region of 

Lake Telets (Altai preserve) is suggested as terra typica res- 

tricta. Nee Cervus sibiricus Schreber, 1784. 
1874. Cervus caspicus. Brook. P.Z.S.L., p. 407. Locations southwest 

of Caspian Sea. 
1876. Cervus eustephanus. Blandford. P.Z.S.L., 1875, p. 637. Tien 

Shan in China. 
1880. Cervus luhdorfii. Bolau. Abh. Naturw. Hamburg., vol. 7, p. 33. 

Northern Manchuria, "Buryatskaya steppe". 
1886. Cervus caspius. Radde. Fauna u. Flora d. sudivestl. Caspi- 

Gebietes, p. 10. Talysh. 
1889. Cervus isubra. Noack. Humboldt, vol. 8, p. 12, fig. 8. Names 

based on the same specimen used for luhdorfii. 
IS92. Elaphus ussuricus. Heude. Mem. Hist. Nat. Emp. Chin., vol. 

2, p. 113. Ussuri. 
1897. Cervus bedfordianus. Lydekker. P.Z.S.L., 1896, p. 932. 

Manchuria. 
1898. Cervus canadensis asiaticus. Lydekker. Deer of All Lands, p. 

104. Region south of Lake Telets near Yenisy source, i.e., the 

present Altai preserve. 
\9QQ. Cervus bactrianus. Lydekker. Ann. Mag. Nat. Hist. ,^vol. 5, p. 

196. Russian Turkestan, probably Bukhara. Also suggested are 

the upper courses of the Amu-Darya and its tributaries in 

Tadzhikistan. 
1902 Cervus wachei. Noack. Zool. Anz., vol. 25, p. 146. Chingil'ta, 

valley, near Chernyi Irtysh, Kobdo. 
\9^Ъ. Cervus vulgaris. Botezat. Morph. Jahrb., vol. 32, p. 154. New 

name for elaphus. 
1903. Cervus vulgaris campestris. Botezat. Morph. Jahrb., vol. 32, p. 

154. Carpathians, Bukovina (lower belt of mountains). 

*"On the title page, 1873; released at the end of 1872. 

*'N.A. Severtsov (1872-1873) states that his material for the description of Tien 
Shan forms was two adult males and one young deer caught "at the end of October in 
Trans-Iliisk Alatau, east of Vernii [Alma-ata], on the peak of Turgena and at Santash 
Pass near the eastern end of Issyk-Kul' " (p. 103). There is no justification for consid- 
ering "Dzhungarsk Tien Shan and probably Kuldja" a type locality (Lydekker, 1915; 
Ellerman and Morrison-Scott, 1951). 



161 

1903. Cervus vulgaris montanus. Botezat. Morph. Jahrb., vol. 32, p. 
155. Carpathians. 

\9M. Cervus hagenbecki, Shitkov. Zool. Jahrb. Syst., vol. 20, fig. 4, 
p. 103. Turkestan. 

1907. Cervus balticus. Matschie. Weidwerk in Wort und Bild., vol. 
16, p. 186. Polish Baltic. Libemyul'. 

1907. Cervus biedermanni. Matschie. Sitzungsber. Ges. Naturf. 
Freunde, Berlin, p. 223. Telets, Altai. 

I9li. Cervus caucasicus. Winans. Amer. Mus. Journ. Zool, vol. 14, p. 
67. Nomen nudum. 

\9\Ъ. Cervus canadensis baicalensis. Lydekker. Cat. Ung. Mamm. 
Brit. Mus., vol. 4, p. 134. Sayan and Baikal mountains west of 
Lake Baikal. Given as substitute for Cervus maral var. sibirica 
Severtsov,* 1872. 

\920. Cervus elaphus brauneri. Charlemagne. Animals of the 
Ukraine. Forests of hilly Crimea. (Formal description not 
available. Only a new name with a reference to the description 
in Brauner's work, 1900 is given). 

1925. Cervus elafus (Sic!) tauricus. Fortunatov. Sbornik Askaniya- 
Nova, p. 324. (name) and p. 326 (description). Crimea and 
Crimean preserve (forest near Koz'modem'yansk monastery). 

1956. Cervus elaphus carpathicus. Tatarinov. Animals of Trans- 
Khidnikh Region of the Ukraine, p. 103. Eastern (Soviet) Car- 
pathians. (V.H.) 

Diagnosis 

Summer coat of adult animals has no spots (a small number of spots 
on the back, sometimes also on the sides, are present as an exception 
in some forms in females, more rarely in males); the first coat of 
fawns is spotted. Speculum large and rises above the croup on the 
base of the tail. Antlers usually have not less than five tines (with an 
ice [bez] tine above the eye besides), which form a crown in some 
individuals. Tail at relatively short; length less than that of ear. 
Height at withers of aduh males exceeds 115 cm. (V.H.) 



♦Spelling of author's name in taxonomic divisions is sometimes at variance writh 
spelling in text and bibliography since the Israeli orthography has been follov^ed in 
this translation. 

"it is possible that this name has an advantage over brauneri since the latter has 
not been assigned fully in accordance with the rules of nomenclature. 

*^This name is a synonym for the forms campestris and montanus. Since their 
descriptions are not completely satisfactory, the name given by K.A. Tatarinov could 
have been of importance to nomenclature but the description has been given without 
conforming to the rules of nomenclature and thus it is essentially a nomen nudum. 



162 

Description 

Large deer; largest species of the genus and the second largest in the 
family. Build slender and proportionate but in general somewhat 
heavier than that of spotted deer. Neck relatively short; head of 
adults fairly raised but relatively lower in fawns. Ears moderate in 
length, proportionate, directed forward, coming up to the eyes and 
shielding them (length about one-half length of head). Tail short, 
roughly about one-half length of ear. 

Antlers of red deer when fully developed usually with not less 
than five tines (two brow, one central, and two terminal). Presence 
of bez tine, which in deer inhabiting the Soviet Union is absent only 
as an exception, is a characteristic feature of the species. In other 
features antlers highly variable, both individually and geographi- 
cally. In several races an increase in number of tines up to 20 occurs 
in extreme cases and even more in some exceptional cases. Individ- 
ual and geographic variability are seen not only in number of tines 
but also in degree of development (size, mass, and weight) of antlers, 
relative size of tines, their position in relation to each other, form of 
beam, and placement of antlers, i.e., degree of proximity of beam 
and position in relation to the sagittal body plane. Antlers may be 
fairly vertical and close-set or turned sideways. Individual variability 
in structure of antlers is fairly high, but each race has a characteristi- 
cally predominant antler type. Moreover, the range of variability of 
antlers in individual subspecies is fairly definite: in some it is slight, 
in others considerable, and in still others maximal. 

In general there are three main types of antlers : 1) Central 
European (hippelaphoid or elaphoid) type. Number of tines large, 
primarily because of branching of antler tips into so-called "flare"; 
tines set like tufts in different planes. Brow, bez, and central tine 
form normally; bases of first two usually close and first or third tine 
best developed. European deer sport such antlers, in which the 
number of tips can be particularly high; this type is also seen on 
Caucasian deer. Also encountered in the Caucasus are antlers of a 
more simple structure, with about five tips not in the form of a 
crown. Thus individual variability is maximum in this type. 

2) Maral type. Antlers larger and can attain extraordinary pro- 
portions. Beam usually thick, with six or seven or more tines in 
exceptional cases. Crowns do not form and terminal tines set in a 



^For more details, see Heptner and Tsalkin (1947). 



163 

single plane (sagittal or close to it). Fourth tine largest in size. Brow, 
bez, central tines well developed; distance between the first two fairly 
125 large. Usually, at the point of deviation of the fourth branch, beam 
bent backward (downward), sometimes very sharply. Antlers gener- 
ally well spread out. This type is characteristic of maral of Tien 
Shan and Siberia and Manchurian wapiti of Siberia and the Far 
East. 

3) Hangul {Central Asiatic) type. Simplest of antler types. Antlers 
relatively small although quite massive and usually with no more 
than five tines (brow, bez, central, and two terminal which form a 
fork). Sometimes a sixth tine occurs but there is no true crown. Fork 
often so set that its plane forms a large angle, sometimes almost 
perpendicular, to the sagittal plane of the body. Bez tine often rela- 
tively high and quite separated from the brow tine; often, central 
tine also set high. Position of the antlers differs but usually they are 
set fairly perpendicular. Deviations are relatively few. This type is 
characteristic of several Central Asiatic forms, e.g., Bukhara deer of 
Middle Asia among Russian forms. 

The color of red deer is characterized by the absence of spots in 
all coats except the first. Yet in some geographic races spots are 
sometimes seen in the summer coat, more often in females than 
males, as an individual variation. When the spots are faint, two rows 
occur along the sides, running along the spine; when the spots are 
more intense, a large number are disposed along the back and sides. 
The frequency of this variation differs in different geographic races. 
Among Russian forms spots are seen in Bukhara deer (C. e. bactria- 
nus) but most often in Caucasian deer (C. e. maral). In other races 
they are imperceptible or extremely rare.^^ Individual coloration, 
primarily overall shade and intensity, is characteristic of all races. 
Thus light rusty and dark brownish-gray are seen among maral as 
extremes, together with all intermediate combinations of these 
shades. Intensity of color of individual body parts and the contrast 
between fairly intensely colored parts, etc., also vary considerably. 

Coloration of deer undergoes intense geographic variability. As a 
typical example the color of Altai maral (C. e. sibiricus Sev.) is des- 
cribed here; the specific characteristics of individual races are dis- 
cussed below under the section "Geographic Variability". Hair 



*'Among forms living outside the Soviet Union spots are particularly frequently 
manifest among Corsican deer (C. e. corsicanus) and evidently also among Atlas deer 
(C. e. barbarus). According to some data spots are fairly distinct in most Corsican deer 
in summer. It is, however, incorrect to assume that the summer as well as winter coat 
is invariably spotted in all adult animals of this form (Flerov, 1952). 



164 



color of males on the back and sides is light and grayish-brownish- 
yellow, varying intensely depending on the predominance of one or 
the other shade. Speculum large, broad, and spreads onto croup: 
coloration of speculum in individual animals varies from dull rust 




124 Fig. 43. Maral (Cervus elaphus sibiricus Sev.). Sketch by Komarov. 



165 

to straw. At the top and from the sides it is rimmed by a dark band. 
The extent of the dark rim varies widely: sometimes it is noticeable 
only from the sides and in other instances also covers the speculum 
from above. The color of this band varies from dull brown to true 
cinnamon. On the neck and back, in the mid-dorsal region, a dark 
streak occurs, which is discernible all along the back, but sometimes 
disappears behind the shoulder blades. On the speculum the conti- 
nuation of the streak running on the back is most often well dev- 
eloped in the form of a fairly bright dark brown band, but this is 
absent in some cases. Tail and speculum of the same color. Legs and 
abdomen dark cinnamon- brown, contrasting sharply with the color 
of the trunk. Mane of long, coarse, cinnamon-brown hair well devel- 
oped on the neck. Head brown or cinnamon-brown but lower lip 
light in color (grayish- or brownish-rust). Rings of a much lighter 
color encircle the eyes. 

The winter coat of females is more uniform in color, without the 
sharp contrasts characteristic of males. Main color of hair on back 
and sides grayish-brown and darker than that of males. No darken- 
ing of abdomen and limbs occurs. Abdomen and inner surface of 
legs lighter in color than sides and back. Limbs grayish or 
cinnamon- brown. Mane on neck much less developed but color 
similar to that of males. Dark streak on back and edge of speculum 
more pronounced than in males. 

It should be pointed out that the color of areas such as the specu- 
lum, its rim, and the dark streak on the back are clearly seen toward 
the end of molt, becoming appreciably duller later. This is equally 
true of males and females. 

The coloration of maral is more uniform in summer than in 
winter, and sex-related color dimorphism is very poor at this time. 
The general shade of body color is brown- cinnamon without dar- 
kening of neck, abdomen, and limbs. The cinnamon shade is some- 
what more vivid in females than in males. The dark streak on the 
neck and back is also more distinct in females although it often 
disappears behind the shoulder blades. The speculum is rust- 
colored, or various shades of this color, but brighter than in the 
winter coat. A dark brown band, sometimes faint, traverses midline 
of the speculum. The dark rim is also brighter than in the winter 
coat. 

Not only the general color tone but also the coloration of indi- 
vidual body parts undergo geographic variability. In different races 
seasonal coloration varies in individual animals and the contrast 
between winter and summer coloration is also variable. In European 
and Caucasian deer the contrast between the relatively bright rusty 



166 

summer coloration and the dull grayish-brown winter coat is sharp; 
it is less marked in maral and Bukhara deer but again prominent in 
izyubr. 

In the first coat of young maral the forehead and crown, upper 
part of the neck, and back and sides of the body are brown and 
relatively dull. The rest of the head, lower surface of the neck, and 
legs and abdomen are lighter in color than the sides and back, and 
the cinnamon color of these parts has an admixture of grayish- 
brown. Speculum large and rusty-red; upper surface of the tail also 
rusty-red. Speculum encircled by a fairly bright cinnamon- brown 
band. A dark brown streak runs along the neck and back, extends to 
the caudal base, and passes through the speculum where it turns 
into a cinnamon-brown color. Along the sides of the streak on the 
back and also partly in the region of the base of the neck two rows of 
distinct, well-defined, white spots occur. These white spots are scat- 
tered on the rest of the back, sides, in the region of the shoulder 
blades, and on most parts of the thighs, where they are also white in 
color but less sharply so and not as close-set. In time the spots 
become less distinct. With the commencement of autumn molt the 
spots gradually disappear and by winter the young animal sports an 
adult coat. 

The coloration of young animals also shows marked individual 
variability, primarily in overall shade and number of spots. In 
related groups coloration of fawns does not differ but differences are 
quite prominent in distinct subspecies. 

In general the intensity of coloration of the young evidently 
corresponds to that of adults (see section "Geographic Variation"). 

The skull structure, typical of the genus, stands out prominently 
due to its large size (differences from sika deer were discussed while 
describing that form). 

The size of body and skull and the gross weight exhibit intense 
individual and geographic variability. Weight of the largest races 
of the 'species exceeds a few times that of the smallest. Furthermore 
weight varies greatly in different seasons. In the period of rut a stag 
may lose up to 20% of its initial weight. The animal attains full 
development by the age of four or five years and, at that time, stags 
of the largest Russian race, maral (C. e. sibiricus), measure as fol- 
127 lows: height at withers 150 to 155 cm, condylobasal length of skull 
up to 450 mm, and weight up to 300 kg. Females are significantly 
smaller and lighter than males. 

Newborn maral measure 74 to 77 cm at the withers and 92 to 101 
cm in body length. Their weight on the first two to six days averages 



167 

16 kg (Tsalkin, 1944).'^ 

These deer sometimes give rise to hybrids with sika. Hybrid char- 
acteristics have been described in the section pertaining to sika deer. 

(V.H.) 

Taxonomy 

Cervus elaphus can be considered the most progressive species not 
only of the genus, but also of the entire subfamily Cervinae. The 
total loss of spots, an infantile feature, in adult animals can be 
considered an advanced characteristic. Far East sika deer (C. nippon) 
should evidently be considered the form nearest to С elaphus. South 
Asian species are more isolated from it. Some are highly specialized. 
(V.H.) 

Geographic Distribution 

These deer are found in forest, forest-steppe, steppe, in part desert, 
and montane regions of the Old World, roughly between 25-30 and 
55-60° N lat. and the very same topographic regions between 35 and 
50-60° N lat. in North America. 

Compared to its original state the geographic range of deer in the 
Old and New World has undergone extreme variations over the 
years. These changes concern general features as well as area, which 
has shrunk extensively. The primary factors affecting range size are 
extermination of the animal, particularly vulnerable because of its 
large size, and land cultivation, primarily in forest and forest-steppe 
regions falling in the temperate zone of the northern hemisphere, 
which represented the original range. 

In some sections of the European part of the USSR these deer 
were exterminated so long ago (probably in the first millenium A.D. 
or possibly at its inception) that it is hardly possible today to recon- 
struct the former habitation of the animal with any degree of 
certitude. 

Geographic Range in the Soviet Union (Reconstructed) 

The boundary of the former range of deer in the European part of 
the USSR encompassed the Baltic region in the east at least up to 
Pskov, but probably more to the east, possibly even up to the south- 
ern coast of Lake Ladoga (encountered in the second half of the first 
millenium B.C. and in the ninth and tenth centuries A.D. at Pskov 



**Some additional data are given in the section "Geographic Variability". For 
further details on morphology, taxonomy, and zoogeography, see Heptner and Tsal- 
kin, 1947. 



169 

and Old Ladoga). However, the animals were very rare. From the 
Baltic the boundary proceeded south, encompassing Belorussia and 
the western parts of the Smolensk region. 

From there the boundary turned east, probably through the 
Kaluzh region (see below). The northern boundary of the range 
between the upper Dnieper and middle Volga is not accurately 
known. Deer undoubtedly lived throughout the Black Sea and Azov 
steppes, even in such places as Perekop, throughout the central 
Caucasian steppes, and also in the steppes around Stalingrad and 
along the Volga up to Astrakhan'. ^^ In the north the occurrence of 
deer is known around Chuguev (close to Khar'kov) in the Belgorad 
region, in the Poltava region, in the regions of Mirgorod, Lubensk, 
Gadyacha, and farther north in Glukhov (east of Chernigov). Deer 
also lived in the Kursk region, Voronezh, and Morshensk (north of 
129 Tambov). In the light of these data one may assume that deer evi- 
dently lived in the Trans-Oka forests (S.V. Kirikov), i.e., in Tul'sk 
and other places of the region south of Moscow. Somewhere south of 
the Oka, or along the Oka, the northern boundary of the range 
evidently proceeded into central Russia during the last 1,000 years or 
even last 500 years. 

Deer also lived around Moscow but at a much earlier period 
(D'yakov culture— 700 B.C. to 500 A.D.). At that time, however, they 
were rare (Formozov, 1947) and evidently exterminated long before 
Moscow was founded, since there are no records of this species. Deer 
which lived in the royal hunting grounds around Moscow (Ismailov 
menagerie, for example) were undoubtedly imported. Therefore, 
there is no adequate basis for considering Moscow an ancestral habi- 
tat of red deer (Flerov, 1952; map).^^ 

Turkin's report (1900), with frequent references to F.K. Lorents, 
about the greatly isolated habitation of deer in the Nikol'sk basin 
district of Vologod, Galichsk-Kostrom, and Semenov-Nizhegorod, 
even in the 1880's and 1890's is plausible but highly dubious. All 
these places are very isolated from the more southern range of deer 

'''information about deer antlers found at Dubovka on the Volga and tfie deer 
hunting of the Kalmyks, according to Pallas (1776), pertains to reindeer and not to 
the species under description as is sometimes stated (Formozov, 1946), although Pal- 
las erred in this case. 

^^That deer were brought into Moscow county not very long ago was first sug- 
gested by Turkin (1900), who saw with F.K. Lorents a relatively freshly shed antler 
found in the 1860's in Bogorod district (Bogorod now named Noginsk). This single 
evidence, even if it is reliable, cannot serve as proof. The possibility is not excluded 
that this antler belonged to an animal escaping that way. In Ismailov menagerie, for 
example, about 700 animals lived even in 1820's and were hunted (Middendorff, 
1867). 



170 

both geographically and chronologically. For the more southern 
regions information is available for a period not later than the eigh- 
teenth or early nineteenth centuries, while finds of deer remains 
belonging to the first half of 1000 A.D. are known from Kaluga 
region (Tsalkin, 1956; for data on Pskov and Ladoga, see above). ^^ 

There is no information whatsoever (except what has been given 
for Semenov basin above) about deer from the area between the 
middle parts of the Tambov and the Urals. Evidently the northern 
boundary of the range, running in mid-Russia somewhere around 
54 to 55°N lat., proceeded eastward at the same level along the 
forest-steppe and farther toward the Urals. Here deer were reported 
on the western slopes not only in the upper reaches of the Sakmara 
(even in the first half of the last century), but also around Ufa (55° N 
lat.; end of eighteenth century, probably still later). The general 
direction of the range boundary of these deer bears a positive anal- 
ogy to the boundary of the range of wild boar and roe deer.'*' 

The extent of dispersal of deer in the Urals is not precisely 
known. Evidently, however, in the southern part of the mountain 
range they were widely distributed and inhabited the environs of 
Sverdlovsk on the eastern slope even in the first half of the last century 
and even in the 1850's (at 57° N lat.). It is highly possibe that west of 
the Urals the northern boundary was not restricted to 55° N lat. 

From roughly 57° N lat. in the Urals the northern boundary of 
the range in western Siberia turned east with a slight southward dip, 
covering Barabinsk steppe from the north, and ran through the 
Mariinsk and Achinsk regions, i.e., roughly along 56° N lat. From 
there, rising slightly north, it turned toward the Yenisey, crossed 
Mendel' River, the left tributary of the upper courses of the Keta, and 
ran toward the mouth of the Angara, i.e., crossed the Yenisey 
roughly around 58° N lat. 

In the region west of the Volga the range extended south up to 
the Black and Azov Seas and included the central Caucasian steppes 
up to the Caspian coast. The range there crossed directly into the 
131 Caucasus where, toward the south, it extended up to the state bound- 
ary. Deer were probably absent in the desert areas on the right 

*'ln Voronezh region, at Ramon', slightly north of Voronezh, deer survived up to 
1917. After this they scattered, when Graf forest was colonized. Later a recent herd of 
deer from Voronezh sanctuary started settling there. One cannot state with certitude 
that Ramon' deer were indigenous forms and not introduced (at least partly). 

As in the case of roe deer, these deer too exhibit significant taxonomic differences 
between the western form, occupying western and central sections of the European 
part of the USSR (European deer), and the form associated with Siberia and the Urals 
(maral). Nevertheless, there is no adequate proof to suggest any effective isolation 
whatsoever between these regions. 



171 

banks of the lower Volga, but on the Volga itself were present up to 
Astrakhan'. 

The southern boundary of the reconstructed range between the 
Volga and Ural Rivers is not known. Based on physical conditions 
and ecological characteristics of the area and by analogy with some 
other ranges, one may only assume that the southern boundary 
probably passed somewhere between Stalingrad or Kamyshin and 
the Ural. There is reason to believe that in old times deer could have 
migrated far south, probably up to the Ural River estuary, along its 
bottomland deciduous forests. 

Beyond the Ural River deer (maral type) were extensively distri- 
buted along the steppes of Kazakhstan, ^^ and encountered along the 
bottomland deciduous forests of river valleys, along some islands of 
forest in the plains, and along some elevations and small hill sec- 
tions. Their distribution there was confined to isolated spots, espe- 
cially farther away to the south. The southern boundary of the range 
evidently covered the Ilek basin and passed via some unknown route 
through the upper course of the Tobol into the Aman-Karagay pine 
forest district (around 100 km south of Kustanay at Semiozernii).^^ 
From there the boundary turned through Kokchetau mountain into 
the Akmolinsk district, covered the distant Karkaralinsk mountains, 
and extended into Chingiz-Tau and then Tarbagatai. It is possible 
that the range boundary ran even more southward and covered the 
pine forest of Naurzum-Karagay (about 200 km south of Kustanay at 
Aksuat), and probably also the Ulutau mountains, and proceeded 
from there into the Karkaralinsk mountains directly to the east, but 
there is no direct proof of this. 

In Middle Asia and southern Kazakhstan'" the range (maral type 
deer) covered Tarbagatai, Saur, Dzhungarsk Alatau, and the entire 
Tien Shan mountain system in the south to the northern fringes of 

"The range and boundaries described are based on data of Middendorff (1867), 
Eversmann (1850), Pallas (1776), Ternovskii (1956), Turkin and Satunin (1900), and 
other sources, mainly from the literature and archive material gathered and processed 
by S.V. Kirikov. 

'^P. Rychkov (1762) wrote that maral is an animal of the steppes. 

"information about hunts by Tamerlane at An-Karagay in 1391 (S.V. Kirikov) 
evidently refers to this place. 

Of the individual finds in Kazakhstan reported by different researchers 
commencing from the eighteenth century, the foUow^ing are mentioned: upper course 
of Ayat (left tributary of the Tobol), Bayanaul'sk, Kokchetau (Zeredinsk, Imantau, 
San-dytkav, Airtau, Maralsinsk, Sinyukhinsk, and Bol'shoi Toktinsk forests), 
Karkaralinsk, Eremdsk (Ermentau), Ken-Kozlak mountains, mountains of Bulga 
(Bugula), and Chingiztau (Kuznetsov, 1948; Antipin 1941; unpublished material of 
S.V. Kirikov). 



173 

the Ferghana Valley and the northeastern extremity of the Alaisk 
range, in the west to Chatkal'sk andTalassk ranges, and in very old 
times probably also Ugam and Kuraminsk. The Trans-Iliisk Alatau 
and Aleksandrov (Kirgizia) ranges constituted the northern rim of 
this part of the range. Information on the penetration of maral 
farther north at the Chu-Iliisk ranges is not available, although it is 
possible that the animal did live there during very old times. It 
penetrated the Pri-Iliisk jungles, however, settled in the Hi estuary, 
and also survived in the Balkhash rushes (Turkin and Satunin, 
1900). In the northwest the range extended up to the Karatau 

75 

range. 
132 Apart from the regions described above there were some habitats 
in Middle Asia occupied by deer of different forms. Some were asso- 
ciated with the Syr-Darya and Amu-Darya. Along the Syr-Darya deer 
(Bukhara or tugai-type deer) dispersed along the tugais of river val- 
leys all along the Aral Sea upward at least to Kzyl-Orda, but 
undoubtedly much higher in bygone times. There is, however, no 
positive information about this. Evidently deer were also distributed 
somewhat east of the river into the desert and probably penetrated 
the lower course of the Sarys and up to Karatau.'^ 

"information about deer in Karatau is reported by N.A. Severtsov (1873) in Kirgizia. 
Some later researchers doubted these data but without proper basis. Severtsov 
conceded that Karatau deer extended into Syr-Darya and suspected that Karatau and 
Syr-Darya deer belonged to a species other than that of the eastern parts of Tien Shan, 
i.e., they were not marals. Insofar as this concerned Syr-Darya, he was of course 
correct; it is impossible to resolve this question for Karatau deer which disappeared 
from there long ago. It is more probable, however, that this form was maral. 

^Severtsov (1873) suggests that the range in Syr-Darya joined the habitat in Kara- 
tau along the extensive shrub lands which existed then; this is highly justified. Data 
for the Sarys were reported on the basis of conjectural and dubious information 
gathered by N.V. Pavlov (1931), although there is nothing improbable in them. The 
form of deer occurring there, whether maral or Bukhara, has also not been estab- 
lished. Either of the two could be assumed with equal justification. 



130 Fig. 45. Reconstructed geographic range of red deer (Cervus elaphus L.) in the 

Soviet Union (scale in km). Question marks indicate uncertainty about the extent of 

occurrence of Cervus elaphus bactrianus Lyd. in the Ural valley, Ulutau district, and 

Syr-Darya valley. Broken section of line indicates suggested northern boundary in 

regions for which there is no information. 

1 — finds of remains at Old Ladoga (beginning of first millenium A.D.); 2 — finds of 
roughly same period around Moscow and dubious information about finds in 1860's; 
3,4, and 5 — extremely dubious, evidently erroneous references about Nikol' basin 
district of Vologod province, Galich-Kostrom and Semenov-Nizhegorod for end of 
nineteenth century; 6, 7, and 8 — erroneous references to Sary-Tash and southwestern 
and northern Ustyurt for 1930's. V.G. Heptner. 



174 

Farther away the range covered almost all of the Kyzylkum 
which, until recently, was extremely rich in saxaul. In winter and 
spring, having moved out of the valleys and deltas of the Amu-Darya 
and Syr-Darya, deer scattered widely not only in those parts of 
deserts which directly adjoin the river valleys and Aral Sea, but also 
in extremely far away places, at least in western and northern parts 
of the desert. It is highly probable that in the growth season of 
ephemeral plants tugai deer from the Amu-Darya delta could move 
out for a small distance into the desert in the adjoining parts of the 
Ustyurt and the Aral Sea coast as they did in northern Kyzylkum, 
where the environment was more favorable to them (saxaul forest). 

Along the Amu-Darya deer moved to wherever reed and tugai 
thickets were available, from the estuary to the source of the river in 
the mountains, i.e., roughly up to 70° E long., to the south or 
southeast of Kulyab. In the upper reaches of the Amu-Darya deer 
dispersed from the river along the tugais, in all of its right-hand 
tributaries, from Shirabad to Kyzyls wherever tugais were present, 
i.e., quite far away during olden times. They were confined, often 
only during some seasons, to interfluves between the tributaries of 
the Amu-Darya and the mountains, especially in Babatag. The pos- 
sibility of deer having reached the foothills of the Gissar range and 
Darvaz mountains has not been excluded although there is no infor- 
mation about this.^' 

It is possible that deer (probably of the tugais type) lived along 
the Murgab but it must have been long ago and there is no positive 
information about this. Deer (probably Caucasian type) lived along 
the Tedzhensk and in Kopet-Dag in the southeastern part in the 
Atrek basin, which penetrated along girkan-type topographic seg- 
ments from adjoining Iranian regions (Caucasian deer).^^ 



Turkin (1900) describes in great detail, citing no references whatsoever, an alto- 
gether unbelievable geographic range for deer in the region south of the Syr-Darya. 
Part of the range described by him evidently pertains to Marco Polo's sheep but his 
references to forests (in eastern Pamir! V.H.) are confusing. 

T'he southern boundary beyond the Volga and the geographic range in Middle 
Asia are based on material from Severtsov, 1873; Kuznetsov, 1948 and 1948a; Shnit- 
nikov, 1936; D.P. Dement'ev, 1938; Bazhanov, 1945; Antipin, 1941; Zarudnyi, 1896; 
Flerov, 1935; Turkin and Satunin, 1900; Heptner, 1949 and 1956; and other sources; 
and also original material of V.I. Chernyshev and V.G. Heptner. 

There is a report (Bazhanov, 1945, based on material gathered through question- 
naires) that deer were observed in the northern (I) part of the Ustyurt desert (1935), in 
the Sarytash (Sary-Tas) region on the Caspian Sea coast deep inside Mangyshlak Gulf 
region (summer of 1940 and winter of 1941-1942) and that, 200 years ago, according to 
the accounts of "old people" (! V.H.), deer were common in the large saxaul forests of 



175 

133 In Trans-Yenisey Siberia and in the Far East the range of deer (in 
the west — maral, and in Trans-Baikal — Manchurian wapiti) revealed 
the following characteristics. Commencing from the mouth of the 
Angara the northern boundary of the range ran along its left bank to 
the mouth of the Chuna (Uda) and along the Chuna southeast 
roughly up to Vydrinsk (Vydrino-Berezovaya). From there, turning 
east, it extended into the Angara approximately in the region of 
Ust-Vihorev and descended southeast into the source of the Lena 
River, roughly toward Kachuga and Anga. From there the bound- 
ary, running for some distance from Baikal and including Baikal 
in the range, turned almost directly northward. North of Baikal it 
encompassed the Chai basin, at least its upper reaches, the Little and 
Great Chui and Vitim (i.e., northern Baikal and Patom foothills) 
and emerged at the Lena slightly east of Mukhtuya at 114° E long. 
From there it proceeded along the left bank of the Lena for some 
distance and reached 61° or even slightly more north (northernmost 
point of habitation of deer in the Soviet Union). Along the left bank 
the boundary extended to a place 100 km above Olekminsk, con- 
tinued to the right bank of the Lena, and proceeded along it to a 
place 100 km below Olekminsk. From there the northern boundary 
of the range sharply descended south through the uppermost reaches 
of the Amga, ran slightly west of Aldansk town, and emerged 
through the Chul'mansk (at the source of Timptonsk) toward Stano- 
voy range. 

The location of the boundary between the above portion of the 
Stanovoy range and the Amur is not altogether clear in some 
respects. Evidently it ran initially along the southern slope of the 
Stanovoy range, especially along the source of the Gilyuya and its 
upper tributaries, and later descending slightly southeast turned 
through the source of the Uda River, upper reaches of the 
Selemdzha, source of the Bureya, and upper reaches of the Amgun, 
intersecting them, to the mouth of the Gorin, entering the Amur 
from the left. On the right bank the boundary passed through the 
mouth of Khelasso River (? V.H.; 51° N lat., i.e., slightly below the 

southwestern U styurt and are at present quite often encountered in the Great Balkhan 
mountam regions. These data are hardly factual and highly incredible. Nor can one 
accept the taxonomic status of these animals, their migration, etc. which the author 
bases on such reports. It can, for example, be demonstrated that over the last few 
decades deer have only very rarely entered Kopet-Dag and the extreme southwest 
at the exact boundary with Iran (one reliable instance is known; Heptner, 1949). Deer 
were obsent in the well-investigated Great Balkhans and there is no positive informa- 
tion whatsoever about their past occurrence. Information about roe deer in the Great 
Balkhans, reported by Bazhanov, is therefore considered erroneous. 



176 

mouth of the Gorin). East of here the range covered parts of the 
basins of Amur tributaries, the Yaya, and Tumnisk, the latter falling 
into Tatarskiy Strait (? — Tumdshi according to Schrenk, 1859). 
Toward the ocean the boundary terminated at De-Kastra Bay (about 
51°40').'^ 

South of the range boundary outlined above deer reached the 
state boundary everywhere and crossed it. Exceptions were the 
steppe and semisteppe zones in southeast Trans-Baikal where this 
species was evidently absent. 

Present Geographic Range in the Soviet Union 

The present range of deer in the Soviet Union (mid-1950's) differs 
sharply from the reconstructed range, the former constituting only a 
small part of the latter. Moreover, instead of a continuous or rela- 
tively continuous range, we now have only one fairly massive zone 
in the east and small sections in other places. Furthermore, at several 
places where dee^^ are now encountered, they are not indigenous; the 
animals were introduced in preserves or game farms under the protec- 
tion of conservation measures far more stringent for deer than for 
other game ungulates. 

In the west deer are often seen in game farms of Latvia (Tsirava, 
Mezhatne, Bukaishi, Valmere, Dene, Vitsgulbene, and some other 
places). These animals were introduced before the First World War 
from Belovezh Forest and later from other places (Germany). In 
Estonia there are a few animals living on the Abruk islands (south- 
ern coast of Sarem Island) and in Audi forest (northwestern Pyarn), 
which are descendants of animals introduced after World War I from 
134 Germany, which died out in other parts during the Second World 
War. Only a few animals have survived in the Audr region (E.V. 
Kumar i; data for 1957). 

In Lithuania deer (of the same origin) are seen on the right side 
of the Nemansk in regions located in the northern parts of the repub- 
lic. Some animals are also present in Kaliningrad region (animals 

The boundary of the range beyond the Yenisey is based on data from Midden- 
dorff, 1867; Maak, 1859; Schrenk, 1859; Gassovskii, 1927; Abramov, 1954; Kopylov, 
1948 and 1950; Podarevskii, 1936; Favorskii, 1936; and unpublished data from D.U. 
Ivanov (Yakutia). 

The sharp deflection of the range boundary from the Stanovoy range to the south 
and the detour of Uda and Amgun basins has been explained by old and recent 
researchers as due to heavy snow in the east. 

These regions number eight: lonishkis (northernmost), Yurbarkas (adjacent to 
Kaliningrad region), Kurshenai (west of Shaulyai), Pakroiis (east of Shaulyai), Tau- 
roge (lower course of Nemansk), Zhagare (near lonishkis), and three regions on the 
left bank of the Nemansk (F.L. Ivanauskas; data for 1957). 



177 

of game farms) but there is no definite information about them. 

In Belorussia deer exist in tfiree areas: Belovezh Forest (between 
Grodno and Brest at 24° E long.), Volozhinsk region (south of Molo- 
dechno), and Ivenets (west of Minsk). Present-day Belovezh deer orig- 
inated from animals introduced from Germany in the middle of the 
last century (Sileziya, 1865); other deer were introduced there even 
later. Deer of Volozhinsk and Ivenets regions are also introduced 
forms (1929), probably from Belovezh (Serzhanin, 1945). 

In the Ukraine the single natural locale of deer is the Carpathi- 
ans. This is quite a large massif associated with forests and high 
altitudes; the area occupied by deer is quite extensive, corresponding 
to the elongated stretch of the mountain range. In the east this seg- 
ment commences in the Putil'sk (Putila) region south of Chernovits 
at the Rumanian border; in the west it runs toward the border with 
Poland and Czechoslovakia between Drogobych and Uzhgorod. The 
boundary of this segment extends from Putila along the Rumanian 
border in the west up to a point south of Rakhov in Tissa.^' Another 
locale of deer in the western Ukraine is situated north of the one 
described above. It is in the form of a belt extending from west to east 
along the border with Belorussia, almost from the Polish border 
roughly from the source of the Pripyat to the Stviga (right-hand 
tributary of the Pripyat, slightly west of 28° E long.). It thus covers 
the region northwest, north, and northeast of Kovel, to the north of 
Lut and north-northeast of Rovno. The southern boundary of this 
belt traverses roughly in a straight line slightly north of Kovel- 
Rafanovka-Sarna, the border between the Ukraine and Belorussia, 
forming the northern boundary in this segment. ^^ The origin of 
these deer is not known but there is no doubt that they are not 
natives but probably represent animals escaping from Polish game 
farms. 

There is a fairly large population of native deer in the forests of 



*'This segment is located in the regions of Volovets, Svalyav, Irshav, Mezhgor, and 
Rakhov of Trans-Carpathian district; Slav, Khyrov, Starosambor, Turkov, and 
Borinsk of Drogobych region; Nadvornyansk, Yaremchansk, Zhabov, DoUnsk, Bolek- 
hov, Peregonsk, Kosov, and Vygod of Stainslav district; and Brodov (Brodov forests) 
of L'vov district (Tatarinov, 1956). 

^^This zone extends to the OHts region of Volynsk district, to Derazhnyansk and 
Sosnov in Rovensk district. It is quite natural to suggest that deer should be encoun- 
tered even in adjoining sections of Belorussia, but no information whatsoever is 
available about this (Serzhanin, 1955). It is possible that they reach the Polish border 
in the west. 



178 

montane Crimea*^ and an altogether isolated colony exists in Voro- 
nezh preserve (forests north of Voronezh), evidently representing 
deer brought from Europe (see footnote 69 above). Small groups of 
deer are present in a wild or semi domesticated state in Askaniya- 
Nova, in the floodplain of the Dnieper at Goloi Pristan and Burku- 
takh (about 35 km east of Goloi Pristan), in Azov-Sivash preserve 
(Biryuchii Island in the Azov Sea, east of Genichesk) and in forests of 
the Pechenezh region (Pechenega) about 50 km southeast of Kha- 
rikov. Deer of no particular breed were brought from Askaniaya- 
Nova to inhabit the last five places and represent a heterogeneous 
population of different forms of species Cervus elaphus (European 
and Crimean deer, maral, and Manchurian and American wapiti) 
and even sika deer. 
135 In Mordov (north of Temnikov at Mokshe), Khoper (on Khoper 
River at Novokhoper), Bashkir (west of Magnitogorsk), and Il'men 
preserves maral were acclimatized.*'* 

Deer occupy a considerable area in the Caucasus. This is a native 
animal existing within the confines of its natural range, is protected, 
including several sanctuaries, which in total area are no larger than 
legally prohibited areas.* Earlier the range was invariably continu- 
ous and extensive and included all the forests of the country; now as 
a result of shrinking forest areas and persecution it has shrunk con- 
siderably and has broken into several fairly large isolated areas. 
Although some increase in the population of deer has occurred in 
the last decade and an enlargement of the area of habitation has been 
observed in some parts of the country, a considerable shrinkage of 
the range compared to that of the early twentieth century is still 
discernible. 

At present the largest segment of deer habitation in the Caucasus 



*^The suggestion that Crimean deer came from animals brought by the Crimean 
Khans is hardly justified. Taxonomically Crimean deer are quite independent and 
occupy an intermediate position between Carpathian (montanus Bot.) and Caucasicin 
(maral Og.) deer. 

'"Maral live in all the listed sanctuaries and have dispersed quite far away from 
Il'men. But their fate in the territory of the now disbanded Bashkir preserve is not 
known; it is possible that they have died. 

In the 1950's especially the mid-fifties, hunting organizations introduced quite a 
large number of deer, mostly maral but partly Askaniya hybrids into quite a number 
of places. Thus maral were introduced into some places of Moscow district and even 
into Estonia. Information about these introductions is very scanty and hence not 
included here. 

•Meaning of passage not clear in Russian original — General Editor. 



179 

falls in the forest regions of the western part of the mountain range, 
i.e., both northern and southern slopes in the area between Kras- 
nodar (about 39° E long.) and almost up to Elburz (about 42°30' E 
long.). Here, in the north, deer almost reach Maikop and, in the 
south, the range boundary extends slightly short of the sea. On the 
whole the outlines of this part of the range form an irregular trian- 
gle with the peak pointing northwest. 

In the east deer occur in the forest zone in the northern foothills 
and the low mountain belt from Nal'chik to Groznyi (46° E long.). 
Some parts of the habitat lie along the Terek on the plains, where 
deer occupy bottomland deciduous forests and Caspian reeds from a 
place slightly below Gudermes. They are also encountered among 
reeds along the lower course of the Sulak and its estuary.*^ 

Deep inside the mountains on the eastern extremity of the moun- 
tain range there are sections of deer habitat in the upper reaches of 
the Avarsk Kois and Samur. There is also an isolated part of the 
range in the forest region facing the Caspian Sea on the low belt of 
mountains between Makhachkala and Derbent, more accurately 
slightly south of Makhachkala and up to the Samur estuary. 

In Trans-Caucasus the range of deer extends as a long narrow 
belt through forests on the southern slopes, approximately from 48° 
E long, on the east to Lagoda (about 46° E long.) on the west. A 
small part of the habitat occurs in the Kur' valley to the southeast of 
Tbilisi, and another of the same size, or slightly larger, north of it, 
and a fairly large one to the west. The last is in the form of an oval 
with the center at Borzhom and is located along Trialet and Imere- 
tinsk mountain ranges. Finally, there is an extremely small section 
southeast of Batumi into which animals probably penetrated from 
Turkey.*^ 

Deer are absent at present in the Urals and in much of western 
Siberia (excluding maral acclimatized in Bashkir and Il'men pre- 
serves). Their range beyond the Urals covers eastern Siberia and the 
Far East and commences in Altai and in regions north of it. In the 
southwest the boundary commences at the state boundary to the 
northeast of Lake МагкакоГ. Running west and northwest it 
137 encompasses the upper Bukhtarma, i.e., southern Altai and Listvyag 
ranges and the southeastern part of Kholzunsk, roughly at the source 



''The jX)sition changed in the mid-1950's due to the drying up of Caspian Sea and 
recession and destruction of reeds. 

''The description of distribution in the European part of the USSR and in the 
Caucasus is based on data from Kalinin'sh, 1950; Serzhanin, 1955; Tatarinov, 1956; 
Korneev, 1952; Vereshchagin, 1947; Heptner and Formozov, 1941; and others. 



181 

of the right-hand tributary of the Bukhtarma-Khamir. From there it 
turns north-northeast and, encompassing the Terektinsk range, 
leaves the Katun somewhere south of Gorno-Altaisk (Oirot-Tura 
and Ulala), probably at the Chemal or slightly north of it. From 
there the boundary runs to the midcourse of the right-hand tributary 
of the lower Biya-Lebed.^^ In the extreme south of Altai the range 
does not reach everywhere directly up to the state boundary; for 
example, the Chuisk steppe is not part of it. 

To the north from the Altai (Lebed') the boundary turns steeply 
north, making large loops and encompassing Salair principality 
and the Kuznetsk Alatau. In the principality the range reaches 
roughly up to 54°30' N lat. and, moving slightly west, its extreme 
western point falls roughly at 85° E long. In the Kuznetsk Alatau the 
range reaches up to 55° N lat. and is bounded by foothills to the east 
(roughly 90° E long.). Beyond approximately 53° N lat. the bound- 
ary turns east sharply, runs along the right bank of the Abakan and 
then the Yenisey, and ascends sharply northward (Yanushevich and 
Balagoveshchenskii, 1952; map). 

The northern boundary of the range east of the Yenisey, i.e., in 
eastern Siberia and the Far East, corresponds to the range described 

^'Detailed information about the geographic range in the Altai has been given by 
Dmitriev (1938) for the 1930's. From that time the range has become somewhat 
enlarged at places, probably quite considerably, i.e., it may be assumed that this 
enlargement did not occur exclusively in the region of the upper Bukhtarma (E.I. 
Strautman, 1953). Nevertheless the range in Altai as outlined by Yanushevich and 
Blagoveshchenskii (1952) appears somewhat exaggerated; the boundary just crosses 
north of Gorno-Altaisk almost at the confluence of the Katun and Biya. Boundary 
descriptions have been compiled from the foregoing references and data collected 
personally by V.G. Heptner. 



136 Fig. 46. Present (mid-1950's) distribution of red deer {Ceruus elaphus L.) in the 

Soviet Union. 

1 — northern boundary of distribution of native deer of central and eastern Siberia and 
the Far East: maral {Ceruus elaphus sibiricus Sev.) and Manchurian wapiti (C e. 
xanthopygos M.E.); and geographic range boundaries of Tien Shan maral (C. e. 
songaricus Sev.), Caucasian deer (C e. maral Gray), Crimean deer (C. e. brauneri 
Charl.), and Carpathian (C. e. monlamis Bot.) (range of Caucasian deer extremely 
generalized and schematic); 2 — locations of natural distribution of Bukhara deer 
(C. e. bactrianus Lyd.); 3 — some places of successful acclimatization of maral in the 
last decade; 4 — location of occurrence of naturalized introduced European deer (C e. 
hippelaphus Erxl.) in Belorussia and in northwestern Ukraine; 5 — approximate 
places of distribution of the same forms in Lithuania, Latvia, Estonia, and Kalinin- 
grad; 6 — some habitations of polyhybrids of Askaniya origin. Arrows indicate places 
of transgression of deer into western Kopet-Dag and southeast of Batumi. V.G. 

Heptner. 



182 

before (see reconstructed range). There is no information about the 
occurrence of deer in this vast territory for the past few centuries. 
The earliest data go back only to the middle of the last century. In 
any case there were no adverse variations of the range boundary. 
Based on stray facts and some general tendencies one may even 
assume, on the contrary, that deer advanced in the last decade even 
farther north in northern Baikal and Yakutia. Only a small section 
of the range in the Far East underwent adverse changes. Now the 
boundary from the upper reaches of the Amgun runs not to the 
mouth of the Gorin but intersects the Amur above, at the mouth of 
the Sungari (right-hand tributary of the Amur) and from there, 
without encompassing the Yaya basin and Tumnin, runs directly 
southeast, emerging toward the ocean at the Samargi River estuary 
(about 47°20'; Abramov, 1954). 

While the entire range boundary and its general outline beyond 
the Yenisey have not undergone major changes, there are considera- 
ble parts of the territory inside the range which are not colonized by 
deer at present. It is not possible to list and describe all of them. 
Attention may be drawn to the sporadic distribution of the animal 
in northern Baikal, in the large zone along the railway track in 
Irkutsk where there are no deer, and some other areas in eastern 
Siberia and the Far East. Throughout this land red deer are distri- 
buted generally less uniformly than roe deer and elk. 
138 In Middle Asia and Kazakhstan the range has undergone 
extremely distinct changes and has shrunk notably. In all the steppe 
zones of Kazakhstan in the east, including Chingiztau, deer have 
disappeared totally. At present they are still encountered in theTar- 
bagatai, Saur, and Dzhungarsk Alatau. 

The present distribution of maral in the Tien Shan is extremely 
limited and difficult to establish. Maral are absent in most parts of 
their former range except for a few places; they are few in number 
and distributed sporadically; at places they are rare and encountered 
singly. Maral live in the Ketmensk range (upper reaches of the Kena, 
Kegensk range) and in the eastern parts of the Kungei-Alatau (Tyup 
River basin). From the upper reaches of this river the boundary of 
maral distribution runs southwest and passes slightly east of Przhe- 
val'sk, encompassing the eastern parts of the Tersk Alatau. From 
there it proceeds in the same direction to the source of the Malyi 
Narym and the upper reaches of the Narym (above the mouth of the 
Malyi Narym) and to the source of the Atbasha, swinging sharply 
southeast later and running toward the Chinese border. This line 
forms the western boundary of the maral range in the Tien Shan 
(A.I. Yanushevich, data for 1957). 



183 

On the Syr-Darya deer are altogether absent at present; on the 
Amu-Darya only a small area is occupied in its midcourse at Darga- 
nat at 41°30' N lat. (former Darganat preserve) and three small iso- 
lated segments in close proximity in the upper reaches of the 
Amu-Darya and along the lower Pyandzha and its tributaries. These 
sections are as follows: 1) along the lowermost reaches of the Kyzyl- 
su from a place northwest of Parkhar up to its mouth, and along the 
Pyandzha from a point east of Parkar below up to the mouth of the 
Kyzyl-su; 2) along the Vakhsha from Dzhilikulya and slightly north 
to the mouth and along the Pyandzha slightly above the mouth of 
the Vakhsha; and 3) along the Amu-Darya valley from a place 
slightly east of the mouth of the Kafirnigan on the Amu-Darya up to 
the boundaries of Tadzhikistan with Uzbekistan (about 67°40' E 
long.) and away from the river (in the north up to Aivadzh) (accord- 
ing to V.I. Chernyshev).** 

Deer were absent along the Murgab in antiquity; they were also 
absent along the Tedzhen, the references of some researchers not- 
withstanding (the latest available information pertains to the 
1880's). Deer could only penetrate occasionally to the Kopet-Dag 
and that, too, only on the extreme border of the range, i.e., extreme 
southwest (V.G. Heptner).^^ 

Geographic Range outside the Soviet Union 

The range outside the Soviet Union (reconstructed) in the Old 
World included Algeria and Tunisia, probably Morocco (informa- 
tion about Senegal-Gambia erroneous), Corsica, Sardinia (evidently 
absent originally in Sicily, Crete, Cyprus, and islands in the Aegean 
Sea), the British Isles, western, central, and southern Europe, and 
southern Scandinavian Peninsula, i.e., in Norway roughly up to 65° 



^*The disappearance of deer along the Amu-Darya and Pyandzha, which rapidly 
continues even now, is associated with the destruction of tugais. Thus, even in the 
1920's and early 1930's tugai deer were dispersed extensively along the lower Pyandzha 
and upper course of Amu-Darya. They inhabited areas from some points at 69°40' E 
long. (Chubek, Moskovskii) to placesslightlyaboveTermez (border of Uzbekistan and 
Tadzhikistan). Along the Kyzyl-su and Yakhsu they reached Kulyab and Bal'dzhuan, 
along the Vakhsha up to Kurgan-Tyube and slightly more north, and along the 
Kafirnigan dispersed north of Shaartuz. Neither along the Surkhandar'e nor in Babatag 
andKaratau (a small elevation at Pyandzha) were they present in those years. The zone 
of habitation along the river was not continuous (Flerov, 1935; V.I. Chernyshev and 
V.G. Heptner). 

^'See p. 174 ff. for further information about the distribution between Caspian Sea 
and Amu-Darya. 



184 

N lat. and in Sweden on the Baltic coast up to 59° N lat. The range 
covers Asia Minor, northern Iran, Afghanistan (evidently only the 
northern part of it), Kashgar (Tarim basin), eastern (Chinese) part of 
the Tien Shan, montane parts of Dzhungaria adjoining the USSR 
and Mongolia, northern Mongolia (Mongolian and Gobi Altai, 
Hobsogol mountains, Khangai, and Khentei), northeastern China 
in the v^est including the Great Khingan, east and southeastern 
China in the south up to Szechwan, Tibet (mainly eastern part), and 
Himalayas including the southern slopes (Kashmir, Sikkim, and 
Bhutan). 

In America the northern boundary of the reconstructed range 
proceeded from St. Law^rence Gulf to the eastern extremity of Lake 
139 Superior and turned northwest roughly to Great Slave Lake (about 
60° N lat.); from there it ran southwest to the coast of the Pacific 
Ocean, roughly 50° N lat. Along the coast the range extended in a 
belt as far as California. 

The southern boundary turned northward in the form of a 
straight line, bypassed Nevada, passed through Utah to the central 
parts of New Mexico, arced northward, and descended as a promon- 
tory into Texas, reaching south to roughly 32° N lat. From there the 
boundary moved northeast roughly into Missouri, descended south- 
east into northern Alabama and Georgia, and departed again north- 
east toward Massachusetts and St. Lawrence Gulf. 

The above reconstructed range no longer exists. Relatively 
important changes occurred in the New World and the once exten- 
sive continuous range has now been reduced to a few isolated sec- 
tions, for example, in Idaho, Montana, Washington, Manitoba, and 
others. Deer in general have survived only as a result of effective 
conservation measures. In view of the fact that extermination of deer 
commenced only from the days of European colonization, i.e., rela- 
tively recently, the outlines of their natural range can be recon- 
structed rather easily. 

In the Old World history has affected the range in a different 
manner. Outside the USSR, namely in Europe, the general bound- 
ary of the range has altered very little. However, distribution, for- 
merly fairly continuous (at the commencement of the Christian era), 
changed long ago. Deer are now found only in some small segments, 
mostly isolated and separated from each other. They exist only in 
game farms as strictly protected and nurtured animals, because they 
represent a favorite object of hunting in Europe (prized for several 
centuries and even now). Many forms of protection, raising, selec- 
tion, and other aspects of game husbandry have been perfected in 
relation to deer in populated areas. All these measures have percepti- 



185 

bly altered the form of European deer over the centuries but none- 
theless preserved them in this part of the range. 

In Asia deer have disappeared completely at several places; in 
some their disappearance took place long ago (eastern and south- 
eastern China) and in others relatively recently, i.e., during the nine- 
teenth century or even in the nineteenth and twentieth centuries. In 
some regions (China, Tibet, and India) deer are not adequately pro- 
tected and their range and population continue to diminish. In 
Tibet deer still survive in pockets only because they happened to 
colonize in sacred forests, public places, monasteries, etc. where 
hunting is forbidden. As in Europe, deer are mainly protected in 
Tibet because they constitute a "feather in the cap"* of human 
conser^vationists. 

Deer have been introduced for acclimatization into many places 
in Argentina, Chile, the USA (Nebraska and Kentucky), Australia, 
and New Zealand. Mainly European deer (Germany, Balkans, and 
British Isles) have been imported, but a few North American ani- 
mals have been included in these experiments. (V.H.) 

Geographic Variation 

Geographic variation among deer is great, the number of forms 
numerous, and the range of characteristics very broad. 

The publication of special monographs (Heptner and Tsalkin, 
1947; Flerov, 1952) notwithstanding, the race structure of deer has 
yet to be satisfactorily explained. Among the large number of forms 
140 described and commonly accepted^*' evidently seven or eight can be 
found in the Soviet Union. They are distinctly separable into three 
groups, viz., western, Siberian, and Middle Asiatic, corresponding 
to the three types of antler forms (elaphoid, maral, and hangul) 
described above. 

Western-Central-European or Elaphoid Group 

1. Central European deer [C. (C.) e. hippelaphus Erxleben, 1792].^' 
Moderate size (height at withers 120 to 125 cm; stags weigh around 
100 to 160 kg). Color of summer coat bright rusty-cinnamon (red 



*Literal translation of Russian phrase is "fellow travellers of the culture" — Sci. Ed. 
'°Heptner and Tsalkin (1947) list 25, Flerov (1952)— 13, Ellerman and Morrison- 
Scott— 18 (1951), and Haltenorth and Trenze (1956)- 15. 

"Synonyms are numerous but none applicable to Russian fauna. 





10 



11 



141 Fig. 47. Types of antlers of some races of red deer (C. elaphus) found in 

the Soviet Union. 

1, 2, and 3— Caucasian deer (C. e. maral); 4, 5, and 6— Tien Shan maral (C. e. 

songaricus); 7, 8, and 9 — Manchurian wapiti (C. e. xanthopygos) from Ussuri 

region; 10 and 11 — Bukhara deer (C. e. bactrianus). Drawing purely schematic. 

From Heptner and Tsalkin (1947). 



187 

deer). Speculum brownish-rusty, rusty, or rusty-white, sometimes 
merging with color on the back. Winter coat dull gray, differing 
intensely from summer coat. Skull with narrow rostral portion; con- 
dylobasal length 375 to 410 mm. Antlers of moderate dimensions; 
number of tines more in this subspecies than in others and form a 
crown. 

The range covers the western European part of the USSR except 
for the Carpathians. Outside the Soviet Union it covers central and 
western Europe (except the Scandinavian Peninsula), England, and 
Spain, and in the east extends up to Sudet[enland]. 

The difference between this form and the nominal Scandinavian 
form is slight and dubious. That the two forms are identical is 
highly probable. 

2. Carpathian deer [C. (C.) e. montanus Botezat, 1903] (syn. cam- 
pestris and carpathicus). Larger than preceding group and approach 
in their dimensions Caucasian deer. Predominant color more gray 
than that of central European deer; antlers very large and tend to be 
simple (many antlers without crowns). 

The range covers the Carpathians. Outside the Soviet Union it 
extends over all of southeastern Europe east of Sudet[enland]. 

The independent nature of this form is dubious and many 
researchers place it as a synonym for the Central European form. 

3. Caucasian deer [C. (C.) e. maral Ogilby, 1840] (syn. caspius, 
caspicus, and caucasicus). Large (height at withers about 135 cm). 
Coloration highly yellowish. Shoulders and thighs dark-colored. 
Sometimes summer coat of adults (females) with spots. Antlers large 
and heavy, sometimes with crown, and sometimes with two-tined 
fork. Number of tines on the average less than in European deer. 
Skull narrow but muzzle broader than that of central European deer. 
Condylobasal length of skull in males 433 to 460 mm; maximum 
skull length 472 mm. 

The range covers the Caucasus. Outside the Soviet Union it 
includes the Caspian provinces of Iran and Asia Minor (?). 

This form occupies a prominently intermediate position 
between European and Siberian races. Some researchers also place 
this form with the nominal race. 

4. Crimean deer [C (C.) e. brauneri Charlemagne, 1920] (syn. tau- 
ricus). Moderate size, intermediate between European and Cauca- 
sian forms, but much smaller than the latter. Maximum skull length 
in males 335 mm (M 405.5) to 454 mm and in females 31 1 to 378 mm; 
condylobasal length of skull in males 320 to 437 mm and in females 

'^See footnote 63 above for significance of names given to Crimean deer. 



188 

302 to 378 mm. Iji the rest of their features Crimean deer are closer to 
Caucasian. Antlers relatively simple, number of tines usually not 
more than six, and crowns rare. In antler structure this form is very 
close to Caucasian deer and, together with it, occupies in this respect 
an intermediate position between western deer and eastern maral. 
The range covers montane Crimea. These deer are absent outside 
the Soviet Union. 

142 Maral Group 

5. Altai maral [C. (C.) e. sibiricus Severtzov, 1872]^^ (syn. asiaticus, 
wachei, biedermanni, and baicalensis). Very large, representing one 
of the largest races of the species. Height of stags at the shoulders up 
to 150 to 155 cm and weight up to 300 kg. Males in winter coat have 
a light gray-brown-yellow color on the sides; neck, shoulders, and 
abdomen darker than the rest of the body and cinnamon-brown. 
Females more uniformly colored and gray- brown. Summer colora- 
tion of both sexes a fairly uniform cinnamon-brown without dark- 
ening of neck, legs, and abdomen. Speculum large, broad, and high; 
descends onto croup; varies in color from dull rust to hay-yellow. 
Antlers very large, with a thick beam, without crown, and usually 
with six or seven tines. Fourth tine largest; at the point of its branch- 
ing off the beam usually curves sharply to the rear (downward). 
Skull broad and muzzle wide. Condylobasal length of skull in males 
reaches 450 mm. 

Newborn deer with body length of 92 to 101 cm, height at withers 
74 to 77 cm, and weight in first week after birth 11 to 22 kg (Tsalkin, 
1944). These fawns are thus larger than those of other deer races in 
the Old World. 

The geographic range covers the Altai, Sayans, and Pri-Baikal 
region. Outside the Soviet Union these deer are found in northwest- 
ern Mongolia. The features of this form resemble those of some 
American forms.^'' 

Some researchers suggest that the name sibiricus Sev., 1872 has been preoccupied 
by the name sibiricus Schreb., 1784 given to reindeer and call the Altai maral asiaticus 
Lyd., 1898 or baicalensis Lyd., 1915. It is on this basis that Ellerman and Morrison- 
Scott (1951) gave the name asiaticus Lyd., 1898 (p. 369) to Altai maral. Yet, as 
explained by these very researchers (p. 376), the name sibiricus Schreber, in general 
has no significance whatsoever from a nomenclatural viewpoint, i.e., even in relation 
to reindeer, and they do not apply it to this species. Although in the situation under 
discussion there is an element of some misunderstanding, it is clear that the use of 
Severtsov's name "sibiricus" is entirely in order. 

Combining this form with the form canadensis of eastern Canada (Flerov, 1952) 
has no basis whatsoever from the viewpoint of taxonomy and is inapplicable and 
incomprehensible zoogeographically. 



189 

6. Semirechian (Tien Shan) maral [C. (C.) e. songaricus Severt- 
zov, 1872] (syn. eustephanus). Large, corresponding in size to Altai 
deer, but with a slightly darker coloration of the trunk and limbs in 
the winter coat. Skull with retro-orbital walls somewhat narrower; 
width of occipital condyles and outer nasal orifice larger than that of 
Altai maral. 

The range covers the Tien Shan and Dzungarsk Alatau. Outside 
the Soviet Union these deer are probably found in the eastern part of 
Tien Shan. 

This is a poorly distinguished form and possibly identical with 
the preceding one. 

7. Izyubr' [C. (C.) e. xanthopygos Milne-Edwards, 1860] (syn. 
luhdorphi, isubra, ussuricus, and bedfordianus). In body size some- 
what smaller than maral and similar to Caucasian deer. Height at 
shoulders 145 cm and weight up to 250 kg; weight may be even more 
in some exceptional cases. Coloration in summer bright reddish-rust 
and in winter gray- brown-yellow. Speculum large and broad, in 
summer reddish-rust and only slightly lighter than the sides, and 
rust-colored in winter. Legs same color as trunk or only slightly 
darker. Antlers quite short and light with a relatively thin beam, set 
more or less straight. Crown absent and tines usually number six but 
sometimes only five. Fourth tine not particularly well developed and 
sometimes smaller than the rest. Muzzle between canines narrower 
than in maral. Condylobasal length of skull 390 to 435 mm. 

The range covers Trans-Baikal and the Far East. Outside the 
Soviet Union these deer are found in northeastern and northern 
China. 

This form is well differentiated. In Trans-Baikal its range 
adjoins that of Altai maral and animals with intermediate features 
have been described. 
143 8. Bukhara deer [C. (C.) e. bactrianus Lydekker, 1900] (syn. 
hagenbecki).^^ Moderate or slightly smaller in size. In this respect 
Bukhara deer are similar to central European deer or even slightly 
smaller. Height at withers about 120 cm. Body coloration light 
yellowish-gray without rusty tones (sandy), being slightly more 
vivid in summer than in winter. Legs light-colored. Speculum well 
developed but approaching in dimensions that of European deer 
(smaller than maral and Manchurian wapiti); upper parts rust- 

'^This form of deer is sometimes known in literature as "hanglu," a popular 
name used for the related (or identical) form in Kashmir. Recently, it has been labeled 
"gavas," "govas," or "gavazd," or even pink (!) deer. There is no justification what- 
soever to alter the very natural and fairly established Russian name and thus clutter 
up the nomenclature with foreign words. ^ 



190 

colored or yellowish and lower parts whitish. Lips and underbelly 
white. Young (in first coat) differ sharply from young of other forms 
in having a light-colored general background. Antlers of medium 
dimensions, usually with five tines. Crown forms very rarely; two 
terminal tines generally bifurcate toward the sagittal plane of body, 
sometimes nearly at a right angle. Muzzle fairly broad. Condylo basal 
length of skull up to 390 mm. 

Newborn fawns 56 to 60 cm high at withers, 78 to 86 cm long, 
and weigh on first day of birth 8 to 9 kg. They are thus significantly 
smaller than maral. Their main shade is much lighter than that of 
maral. 

The range covers Turkestan. Outside the Soviet Union these 
forms are found in northern Afghanistan. 

This subspecies differs very sharply from all other Russian 
forms. Its relation to the Kashmiri form С e. hanglu, and through it 
to the Himalayan C. e. af finis needs to be accurately established. 
Closeness to the Kashmiri form is indisputable. 

Outside the Soviet Union the following forms of the species are 
acknowledged: С e. barbarus Benn., 1837 (northwest Africa); С e. 
corsicanus Erxl., 1777 (Corsica); С e. hispanicus Hilz., 1909 (Iberian 
Peninsula); C. e. atlanticus Lonnb., 1906 (Norway); C. e. scoticus 
Lonnb., 1906 (Scotland); С e. elaphus Linn., 1758 (southern 
Sweden); C. e. ajjinis Hodgs., 1841 (eastern Himalayas and Tibet); 
C. e. hanglu Wagn., 1844 (Kashmir, probably identical to preceding 
form); С e. wallichi Cuv., 1812 (southeast Tibet); C. e. macneilli 
Lyd., 1909 (central China, Szechwan); C. e. alashanicus Bobr. and 
Flerov, 1935 (Alashan, southeast Mongolia, northern Kansu, and 
Shansi); С e. merriami Nels., 1902 (Arizona and New Mexico, USA); 
С e. nannodes Merr., 1905 (California, USA); and С e. canadensis 
Erxl., 1777 (rest of range in North America). The number of Asiatic 
and probably European forms in this list of Russian forms will no 
doubt be reduced in the future. (V.H.) 

Biology 

Population. The present-day geographic range of red deer in the 
Soviet Union is about one-half smaller than it was a few centuries 
ago. Deer are presently found in considerable numbers mainly in 
preserves and in a few regions in eastern Siberia and the Far East. 
The current Baltic population of deer developed from animals 
acclimatized at different periods. In Lithuanian Soviet Socialist 
Republic in 1952 there were about 200 deer (Kuznetsov, 1954). This 
number had risen to 280 by 1955 (Ivanauskas, 1957) and to not less 



191 




Fig. 48. Crimean deer. July, 1955. Photograph by P. A. Yanushko. 



than 300 by 1957 (F.L. Ivanauskas). At present, on the whole, the 
deer population in the Baltic region is not less than 400 to 450 
animals. 

In Belorussia deer are common only in the preserves of Belovezh 
Forest (72,000 hectares), where their 1953 population was 565 (684 in 
1950 and 771 in 1951; Bannikov and Lebedeva, 1956). Until World 
War I, when Belovezh territory comprised 129,000 hectares (110,000 
hectares covered with forests), deer were far more numerous — 5,000 
to 7,000 animals or more m 1907 through 1914 (Wroblewskii, 1912; 
Severtsov, 1940; and others), or 45 to 64 animals per 1,000 hectares of 
forest-covered area. These deer originated from animals introduced 
from western Europe. 

In the Ukraine red deer are now most abundant in Trans- 
Carpathia followed by the Stanislav region; still their population as 
a whole in the republic (excluding Crimea) barely exceeds 1,000 to 
1,200 animals.^'' Both native and introduced deer dwell here.^^ 



'^Including over 300 in Biryuchii Island in Azovo-Sivashsk preserve. These are 
hybrid Askaniya deer. 

''For more details on acclimatization of deer, see the section below entitled 
"Economic Importance". -- 



192 

In Crimean preserve (30,000 hectares) in 1950 there were 2,096 
deer or over 70 animals for every 1,000 hectares of land. In the rest of 
Crimea deer numbered 1,093 (hence, for Crimea as a whole, the total 
was 3,200), with density in different regions ranging from 0.7 to 16 
per 1,000 hectares (highest in Alushtinsk and Kuibyshev regions) 
with an average of about 5.0 animals (Yanushko, 1957). Subse- 
quently hunting deer was permitted in Crimean preserve and their 
population decreased. Then hunting was banned and the popula- 
tion increased to 2,224 animals in 1957 (A.A. Tkachenko). 

In the rest of the European part of the USSR, excluding the 
Caucasus, deer are found in significant numbers only in the Voro- 
nezh region, where their population in 1952 exceeded 800 animals 
(Mertts, 1953). By 1955 the number of deer had risen to about 880 
animals in Voronezh preserve alone (31,000 hectares), i.e., about 28 
animals for every 1,000 hectares of land (Zharkov, 1957). Due to a 
drop in numbers in the severe winter of 1955 to 1956 and catching 
for purposes of acclimatization, the number of deer decreased (648 
animals were counted in 1956 in Voronezh preserve, 448 in 1957, and 
496 in 1958; P.P. Kaznevskii). 

In the Caucasus the maximum number of deer formerly lived in 
Caucasian preserve. In 1939, when the territory of the preserve com- 
145 prised about 300,000 hectares, there were up to 2,880 deer in it (Zhar- 
kov, 1940). Following a reduction in the area of the sanctuary in 
1951, conditions for deer worsened and, in subsequent years, their 
number in western Caucasus decreased. In Trans-Caucasus the max- 
imum number of deer occurred in Trans-Katal and Lagoda preserves 
(over 1,000 animals in all) and in the territory of Borzhom preserve 
(there were 1,404 deer in 1943; Z.S. Ekvtimishvili; now probably 
less). 

A small population of maral live in the southern Ural. Maral 
were introduced there in Bashkir preserve in 1940 to 1941; in 1951 
they numbered about 350 (Kaznevskii, 1956). 

Accurate figures are not available about the deer population of 
Siberia. They are found in maximum numbers in Altai preserve and 
are quite numerous at some places in the Sayans and Tuva Auto- 
nomous region. Until very recently they were common in the 
Baikal-Lena watershed region, in the upper course of the Lena and 
Kirenga, and at places along the left bank of the Angara. Deer were 
numerous in the Badaibinsk region of Irkutsk (Skalon, 1951) and 
quite common in Southern Yakutia where their population has 
even slightly increased in recent years (Belyk, 1953). 

In the Far East the maximum number of deer occurred in 
Sikhote-Alin: their population was estimated as 10,000 in 1940 in 






:$ 






.""*s. 



X*'M<*k-* -Л 















*AJ. 














194 

Sikhote-Alin preserve (1,700,000 hectares) (Kaplanov, 1948). Follow- 
ing a reduction in the area of the preserve in 1951 the deer popula- 
tion decreased sharply. 

Bukhara deer and maral are distributed in the territory of the 
Middle Asian republics. The maral population in Middle Asia does 
not exceed a few hundred. Bukhara deer have been protected mainly 
in the Amu-Darya basin — up to 100 animals in Darganatinsk tugais 
(Klyushkin, 1954; Dement'ev et al., 1956) and several hundred in 
Tigrovaya Balka preserve (23,000 hectares). In 1946 in the latter area 
138 animals were counted and in 1949—222. Their number has 
recently increased to not less than 500 to 600 animals (Chernyshev, 
1958). 

On the whole the population of red deer in the Soviet Union 
comprises several tens of thousands of animals. 

The former abundance of deer can be assessed with some caution 
from herd sizes, which are more or less proportionate to the total 
population and their saturation of the land. In Laba and Beloe 
basins, where the Caucasian preserve was later established, Dinnik 
(1910) sighted from the end of the nineteenth to the early twentieth 
century herds of 26 (Umpyr'), 35 or 36 (Bol'shoi Bambak), and 70 to 
75 (Urushtensk) females; once he saw a herd of 13 males. Hunters 
have reported herds of 40 to 50 animals. ^* Over a five-year period, 
from 1933 to 1937, in Caucasian preserve only once were 30 deer 
sighted (in summer) in the upper course of the Kholodnaya; 20 of 
these (15 females and 5 juveniles) formed a single herd (A.A. 
Nasimovich). 

Habitat. Their high ecological flexibility enables deer to survive 
under diverse conditions in nature from sea coasts to alpine or 
mountain tundra belts (in the Dzhungarsk Alatau mountains up to 
3,000 m; Shnitnikov, 1936). The range of this species includes taiga 
and broad-leaved forest zones, steppes, and deserts (in semideserts 
and deserts, mainly in tugais along the banks of reservoirs and sax- 
aul bushes), and also subtropical areas in the southern Soviet 
Union.'^ Thus deer inhabit not only regions with an extremely arid 
climate and snowless winters, but also areas with abundant rainfall 
and heavy snow in winter. 

The extreme ecological plasticity of deer can also be judged from 
the results of successful acclimatization experiments with hybrid 



'* In Crimean preserve herds of 40 animals are sighted even now (Yanushko, 1957). 
In subtropical areas (south of Trans-Caucasus), as also in the steppes, deer at 
present are negligible. 



195 



146 forms (obtained by crossing Altai maral with Crimean deer having 
the blood of other forms) on Biryuchii Island in Azov-Sivash pre- 
serve, which reveals that they can survive even in the absence of fresh 
water (M.D. Shpigov). 

For most of the year deer avoid evergreen and dense forest masses, 
preferring more sparse sections with an abundance of deciduous 
trees, shrubs, and herbaceous vegetation, and interspersed with mea- 
dows and glades. Especially convenient for deer life are places with a 
great variety of habitats. In the past, evidently, unusually large 
numbers of deer were encountered in the forest-steppe. Depending 
on the extent of winter snow deer may migrate en masse into the 
forest in winter (in places where the winter is more snowy), or on the 
contrary congregate in the steppes (regions with a continental 
climate). 

At present the main deer reserves are concentrated in montane 
regions since these animals have become extinct on the plains or 
have disappeared as a result of topographic changes. In the moun- 
tains deer inhabit diverse habitats and often are encountered on 
steep and highly rugged slopes which elk usually do not reach. In 
general, wherever red deer, elk, and roe deer are found in the same 




Fig. 49. Maral in spring on thawed patches. Bol'shoi Bashort range. Bashkir 
preserve. April, 1949. Photograph by P.F. Kaznevskii. 



196 

locality, red deer usually utilize a much broader range of habitats 
than the other ungulates. 

Of special importance to deer life in the European part of the 
range are the following: in the Carpathians — beech forests from 
where deer often climb up to mountain meadows or glades (Tata- 
rinov, 1956); in Belovezh Forest — abundant deciduous forests with 
undergrowth and herbage consisting of young pine and mature 
pine-fir-oak forests, or even alder forests in dry years (Sablina, 1955); 
and in Voronezh preserve — mixed and oak forests, and pine forests 
in winter if there are abundant shrubs and a leafy undergrowth 
(Mertts, 1951). In Crimean preserve in summer a maximum number 
of deer are seen in oak forests, followed by beech; in winter up to 95% 
of all deer sighted were located in oak forests (Yanushko, 1957). In 
the southern Urals maral acclimatized there are attracted to hilly 
pine-larch forest steppes and avoid evergreen, humid, and deciduous 
forests (Kaznevskii, 1956). 
147 In Caucasian forests deer occur from the coastal rushes of the 
Caspian Sea (Dinnik, 1910; Heptner and Formozov, 1941) to high 
alpine meadows inclusive. In summer in Caucasian preserve deer are 
particularly numerous in the upper one-third of the forest belt 
(glades with tall grass vegetation, a zone representing the upper 
forest limit) and in subalpine and alpine meadows (up to 2,500 m). 

In the Tien Shan mountains the main habitations of maral are 
fir forests interspersed with glades (Shnitnikov, 1936; Kuznetsov, 
1948; and others). In southern Siberian maral and Manchurian 
wapiti use diverse habitats. Of special importance are burned-over 
forests with growing leafy seedlings and abundant grass. In mon- 
tane regions the maximum number of deer are seen in the central 
parts of the forest belt of the mountains in the Altai and Sayans, and 
in the subalpine zone in summer from where they go even into 
alpine meadows. In the Sikhote-Alin mountains Manchurian wapiti 
abound in moderately old burned-over forests of the Manchurian and 
Okhotsk type and also in coastal oak forests. They are considerably 
fewer in coniferous forests. In summer some animals ascend to mon- 
tane meadows at a height of 1,700 m (Kaplanov, 1948). 

In montane regions (Caucasus, Altai, Sayans, and Sikhote-Alin) 
adult males are confined to the mountain slopes at slightly higher 
altitudes than females and young. However, in Caucasian preserve, 
in some habitats in alpine meadows, mostly females and young are 
seen in summer (upper courses of Kholondnaya) as well as in winter 
(Pshekish range) (Nasimovich, 1936 and 1955). 

Extremely typical habitats of Bukhara deer are now mainly found 
in tugais of Amu-Darya basin. In Darganatinsk tugais deer are con- 



197 

fined to prickly shrubs of the salt tree {Halimodendron holoden- 
dron), along sections with luxurious reforestations of poplar 
(Populus pruinosa), in bushes of oleaster [Eliagnus angustifolius) 
and in reed thickets. In summer deer are usually seen in association 
with an abundance of dense and high herbaceous vegetation and are 
frequently encountered in autumn and winter among forest planta- 
tions (Klyushkin, 1949 and 1954; and others). Along the Kafirnigan 
River deer inhabit a narrow belt of reed thickets with sparse bushes 
of oleaster. Along the Pyandzha and the lower reaches of the Vakhsh 
they occur mainly in flooded forests of swamp poplar mixed with 
oleaster. Particularly favored sections are small clearings with her- 
baceous vegetation and reed thickets (Bobrirlskii, 1933; Flerov, 1935 
and 1952; and others). In the cold season of the year Bukhara deer 
penetrate adjoining deserts, where they live in saxaul and tamarisk 
shrubs, avoiding extremely exposed and hilly areas (Flerov, 1935; 
and others). In the past they were found much more regularly in 
desert regions (Bobrinskii, 1933; Antipin, 1941; Bazhanov, 1945; and 
others). Migrations from tugais into adjoining deserts are also com- 
mon in early spring, when ephemerals begin to emerge. 

Bukhara deer differ notably from all other subspecies of red deer 
occurring in the Soviet Union, not only in relation to habitat but 
also in all other biological characteristics. These differences are 
much more significant than, for example, the corresponding differ- 
ences between various species of turs. 

Food. The food of red deer has been studied mainly in preserves; 
in general, no less than 300 plant species (including fungi) are con- 
sumed by deer, but detailed data have not been published so far. For 
Bashkir preserve over 150 plant species have been established as food 
plants for deer (Kaznevskii, 1954),'°° around 150 for Voronezh pre- 
serve (Mertts, 1953; and others), '°' over 130 for Crimea (Yanushko, 
1957), not less than 80 for Belovezh Forest (Sablina, 1955; and 
others), over 70 for the Far East (Abramov, 1954),'°^ about 60 for Cauca- 
sian preserve (Nasimovich, 1939 and 1940), over 50 for Altai preserve 
(Dmitriev, 1938), and up to 50 for Azov-Sivash preserve (V.A. Matts- 
kov; M.D. Shpigov; Ishunin, 1956; and others). Data are even more 



""^ According to Danilov (1958) deer use in their food 234 types of angiosperms alone 
(61 types of trees and shrubs and 173 types of herbaceous plants). 

""The full list of food plants has not been published. 

'"^The full list of food plants has not been published. Kaplanov (1948) has listed 
slightly more than 40 plant species consumed by Manchurian wapiti in Sikhote-Alin 
preserve. 



198 

scanty or altogether fragmentary for other regions. 

Of utmost importance in the plant food of deer are edible por- 
tions of trees and shrubs (branches, shoots, bark, leaves, buds, and 
needles of conifers) and fruits of some trees (beech, oak, etc.). In 
winter lichens are also consumed in some areas and horsetail in the 
Far East. Seedlings of reeds and saltworts are avidly eaten by Buk- 
hara deer. Depending on the nature of the vegetation, extent of snow 
in winter, and numerical strength of ungulates, the frequency of 
herbaceous, tree, and shrub (branches) intake varies greatly. 

Tree and shrub food are of great importance year-round, includ- 
ing summer, for deer in Belovezh Forest and especially in the Cri- 
mea. Thus in Crimean preserve food plants include 94 species of 
herbaceous plants and 37 trees and shrubs (81 and 27 respectively in 
summer). In the stomachs of six deer caught between May 27 and 
August 10 were found the remains of leaves, buds, and young shoots 
of trees and shrubs accounting for 60 to 70% of the total weight of 
food. In autumn and winter the remains of tree and shrub foods in 
the stomachs of deer comprise 60 to 85% of the total food mass 
(Yanushko, 1957). 

Herbaceous, tree, and shrub food are of equal importance in the 
diet of deer in Voronezh, Bashkir, and Caucasian preserves. In 
summer mainly herbaceous vegetation is consumed. In Voronezh 
preserve tree and shrub vegetation and dried plant parts are mainly 
used in winter. "^^ The food regime is roughly similar in maral of 
Altai preserve where herbaceous vegetation enjoys predominance in 
summer; in winter tree and shrub vegetation and dried plant parts 
are consumed to an equal extent. 

In the diet of eastern Siberian deer, especially at places where 
winters are not extremely snowy (eastern Sayan, central Baikal, and 
Chita River basin), herbaceous vegetation is more important and 
deer consume it in considerable quantities not only in the snowless 
period, but also in winter, mainly in the form of dried plant parts 
(Pavlov, 1949; Kopylov, 1950; and others). Herbaceous vegetation is 
also extremely important in the diet of Far East Manchurian wapiti 
(Kaplanov, 1948; Abramov, 1954). 

Depending on the season of the year and the availability of food 
resources, Bukhara deer consume fairly large quantities of herbace- 
ous plants (wormwood [sagebrush] very avidly) or trees and shrubs 
(shoots of saxaul and tamarisk, poplar leaves, bark of young 
poplars, oleaster, and others) (Flerov, 1935; and others). In Dargana- 

In Caucasian and Voronezh preserves fruits of nut-bearing trees are of vital 
importance in tfie winter diet of deer. 



199 

tinsk tugais the most favorite and universal food of deer is the tender 
and succulent leaves of poplar {Populus pruinosa). These leaves are 
well consumed in summer as well as in autumn and winter when 
they shrivel and drop. Bukhara deer avidly take to the halophyte 
Aeluropus Utoralis and, in early spring, ephemerals, especially desert 
sedge (Carex pachystilis) (Klyushkin, 1949). In the tugais of Tad- 
zhikistan, Bukhara deer consume, apart from herbaceous foods, large 
quantities of tamarisk, saxaul, and saltwort in summer which, by 
autumn, become the main food; in winter, fruits of oleaster acquire 
utmost importance (Chernyshev, 1958). Succulent rootstock, bulbs, 
and other underground plant parts may play an important role in 
the ration of Bukhara deer, but this aspect has not been adequately 
studied. 

Azov-Sivash hybrid deer, because of sparse tree and shrub vegeta- 
tion in the region, feed almost exclusively on herbaceous plants 
(legumes and grasses are particularly important). Often they eat 
common reed {Phraginites communis) — its tender shoots, succulent 
149 and sweetish underground root stock, and tender flowering panicles 
still in their sheaths (M.D. Shpigov). In autumn roots of sea kale 
(Cram.be pontica) and Seseli are avidly consumed; deer dig them up 
from the soil with their hooves (G.I. Ishunin). 

One-eighth to one-fourth of the plant species consumed by deer 
can be considered the primary food taken in substantial quantity. 
For example, in Crimean preserve, of the 108 plant species used by 
deer in summer, 21 species are eaten avidly and often, 32 species to a 
lesser extent, and 55 species rather rarely (Yanushko, 1957). 

Among herbaceous plants grasses are consumed regularly (their 
importance is particularly great in spring and early summer before 
their coarsening; they are also consumed in winter in large 
amounts). Other plants consumed avidly are thqse of families Com- 
positae (including wormwood), Leguminosae, and Umbelliferae. 
When food is abundantly available, the most nutritive or succulent 
parts of herbaceous plants are used in the first instance; in the latter 
half of summer flowers and the upper parts of stalks are avidly eaten 
(western Caucasus, Belovezh Forest, etc.), and in winter the root 
portions of green stalks and the leaves near the roots of perennials. 
Grasses which remain green in winter (mountain fescue and others) 
represent the favorite food of deer in the Caucasus, Altai, and the 
Sayans. Deer eat good hay more readily than twiggy food. In spring 
forest squill, pasque flower, and some other early spring plants are 
consumed well. In Siberia a particularly favorite deer food in 
summer is buckbean; in northern Mongolia leeks are avidly con- 
sumed in autumn (Bannikov, 1954). Deer also eat belladonna, aco- 



200 

nite, and other poisonous plants well. 

From among trees and shrubs deer in general use no less than 70 
species; of particularly great importance are oak, some maples, elm, 
ash, hornbeam, witch elm, beech, aspen, birch (Caucasus and east- 
ern Siberia), lime, pear, rowan, bird cherry, willow, prickwood, 
hazel (mainly catkins), blackberry, raspberry, jasmine. Guelder rose, 
buckthorn, currant, honey-suckle, ivy, mistletoe, and many other 
species at different places. For example, in the Far East deer con- 
sume well, among others, Amur philodendron, Manchurian aralia, 
and lespedeza; in the Caucasus yew (upper portions of small trees); 
in Crimea smoke tree {Rhus cotinus) and others. When other food is 
not abundantly available in winter, pine shoots with needles and 
even juniper are eaten in significant quantities (Belovezh Forest, 
Mordov, Voronezh, and Crimean preserve). In the Caucasus green 
leaves and stalks of ephemerals represent a main winter food item; 
ivy leaves are also very important in winter in Trans-Caucasus. In 
spring flowers of Dahurian rhododendron are also eaten well (Gas- 
sovskii, 1927). Fir, spruce, and larch are eaten rarely and that too 
only when there is a paucity of food in winter. Excessive consump- 
tion of pine needles, a food deer may be forced to eat, can disturb the 
functioning of the gastrointestinal tract and cause poisoning among 
young animals (Mertts, 1953). '"'* In Belovezh Forest instances of 
death of deer due to excessive intake of spruce needles have been 
recorded (Wroblewski, 1927). 

Depending on abundance, accessibility, preponderance, and 
chemical composition (which has never been studied), the impor- 
tance of a particular species of tree as deer food can vary considera- 
bly. For example, the branches, shoots, and dry leaves of oak serve as 
the major winter food of deer in Crimean preserve; oak in this region 
is consumed more often than any other tree or shrub (Yanushko, 
1957). In Belovezh Forest oak occupies only fifth place after other 
tree species, while in Voronezh preserve it drops to tenth place (Kaz- 
nevskii, 1958). 

In Bashkir preserve lime and juniper are fairly common but deer 
do not consume them; yet in Belovezh Forest lime is eaten quite 
150 often and even juniper is consumed (P.F. Kaznevskii). In places 
where winter is relatively mild and thaws frequent (southwestern 
USSR, Crimea, the Caucasus, and Primor'e region) not only 
branches, shoots, etc., but also soft bark are eaten; in general, how- 
ever, the importance of bark is particularly great in transitional 

In the snowy winter of 1955 to 1956 deer in Voronezh preserve consumed pine 
needles in large quantities (Kaznevskii, 1958). 



201 

seasons, i.e., in autumn and later in spring. In rare cases deer also eat 
bark of coniferous trees, for example pine and more rarely spruce, 
along with bark of deciduous trees. Pine bark is eaten in small 
quantities in summer when deer also gnaw at the upper root por- 
tions of spruce (Latvia; Kalnin'sh, 1950). 

In the latter half of summer and in autumn deer avidly consume 
fruits of many fruit-bearing trees; Damson plum, pear, apple, and 
others; they also consume the succulent fruits of some shrubs as well 
as berries. Acorns and beechnuts are particularly important in the 
Caucasus. Deer utilize acorns as food not only in autumn but also in 
winter; when the snow is not more than 25 to 30 cm thick, it is not 
difficult to dig them up with muzzle and hooves. They also eat hazel 
nut, chestnut, pine nuts (Baikov, 1915; Yanushevich and Blagovesh- 
chenskii, 1952), lime, etc., but these food sources, compared to 
acorns and beechnuts, are not very important. 

Deer consume many types of mushrooms including white and 
peppered, russula, honey agaric, etc. In winter, especially in snowy 
periods, deer often consume epiphytic lichens {Usnea barbata, Par- 
melia, and others). Red deer have been observed time and again 
digging under snow for lichens (down to 20 to 25 cm) just as rein- 
deer do when there is a food shortage. Lichens dug up include 
Cladonia alpestris, C. rangiferina, C. sylvatica, Cetraria islandica, 
and others (Belovezh Forest, Voronezh, Mordov, Azov-Sivash, and 
Crimean preserves, and northwestern Trans-Baikal). This pheno- 
menon underscores the extraordinary ecological adaptabilty of red 
deer. It has also been recorded that deer consume both fresh- water 
(filamentous) and marine algae washed ashore. The consumption of 
marine algae and other sea products washed up on coasts (crabs, 
fish, etc.) is evidently motivated by the animal's need to meet min- 
eral and possibly protein requirements. The volume of food in the 
stomach of a deer reaches 30 liters or more (April; Mertts, 1954). 

Over much of the range deer visit waterlogged and sometimes dry 
salt licks. In winter deer clamber onto ice crusts on rivers and lick 
ice, which is rich in mineral matter because formed from montane 
water (Kirilov, 1902; Kaplanov, 1948; Okhotina, 1957; and others). 
When a shortage of fresh water occurs, deer drink sea water, taking it 
in small gulps (Azov-Sivash preserve; V.A. Mattskov; and others). 
Drinking sea water has also been observed in Primor'e region 
(Kaplanov, 1948) and probably follows the same pattern as in salt 
licks. Bukhara deer lick salt outcrops (Flerov, 1952). Visits to salt 
licks are most frequent in spring and the first half of summer; such 
visits also occur during the period of estrus but are far less frequent. 

Home range. Fairly fixed home ranges are quite characteristic in 



202 

those sections of the range where the population lives year-round in 
the same place, without resorting to major seasonal migrations 
(Graf, 1956). In the Soviet Union this is observed, for example, in 
Voronezh preserve, where deer are relatively settled and territorial 
behavior is relatively well manifested. A complete change of biotope 
and significant changes in herd characteristics are typical of the 
transitional season; both are highly constant throughout the rest of 
the year. Adult females with fawns or at the head of a herd often shift 
home range in summer (Graf, 1956); adult males typically relocate 
with harems in the period of rut (Mertts, 1953; Graf, 1956). 

Mention is made in literature that deer mark the boundaries of 
their territory by various methods ("loners" and also adults acting as 
leaders of herds). Such territorial demarcations include "scratches" 
at definite places on the bark of trees and odoriferous deposits, due 
to rubbing up against trees, urination, etc. Animals not belonging 
to a given herd and entering a previously claimed territory are usu- 
ally driven away (Graf, 1956). Territories are generally separated by 
151 neutral zones, but if the population density is high, as in Crimean 
preserve, the boundaries overlap (territory of loners and herds) (P.A. 
Yanushko).'°^ In the course of a day deer use only a part of the 
territory. In visiting salt licks they go beyond the boundaries of their 
territories, sometimes up to 4 or 5 km; salt lick zones are therefore 
neutral. 

The size of the home range depends on the availability of food in 
the area and the population density. In winter, when places suitable 
for supporting deer life are usually smaller than in summer, animals 
congregate in a limited territory. '°^ Size is minimal in the latter half 
of winter, especially if it is snowy. In summer in Voronezh preserve a 
small herd of deer is confined to a segment of 300 to 400 hectares, 
while in winter even a herd of 10 animals moves within a territory of 
not more than 200 hectares. With heavy snows a deer herd often will 
not wander more than a kilometer. In the period of fattening, beds 
are set up at intervals of 150 to 200 m (Mertts, 1953 and 1957; and 
others). Maral acclimatized in Mordov preserve traversed 2,200 to 



'"'Black-tailed mule deer {Odocoileus hemionus) (California, USA) in such an 
event vigorously defend only the central part of their home range (Dasman et al., 1956). 

'°% Altai (Dmitriev, 1938), Caucasian (Zharkov, 1940), and Voronezh (Mertts, 
1953) preserves the total area of the main wintering sites of deer constitutes only 30 to 
40% of the area of their summer habitats. In some wintering areas deer density reaches 
40 to 50 animals per 1,000 hectares (Caucasian preserve; Zharkov, 1940). In Voronezh 
preserve, toward the end of the snowy winter of 1955 to 1956 it even exceeded 60 (P.F. 
Kaznevskii), thinning down by five to eight times in summer. 



203 

2,400 m every day on a loose snow cover of 40 to 55 cm depth in 
February-March (F.D. Shaposhnikov). In Bashkir preserve, where 
maral are also acclimatized, their daily wandering in different sea- 
sons varies from 200 to 300 m (in winter when feeding on hay) to 4 to 
5 km. In summer some females, from fawning to estrus, live in a 
sector of 90 to 140 hectares, traversing daily not more than 500 to 700 
m (Kaznevskii, 1956; and others), fn Crimean preserve, where deer 
density is exceptionally high (see above) and trees and shrubs have 
been highly denuded, a home range is limited to 40 to 140 hectares, 
on the average 70 hectares, and daily wanderings to 4 to 5 km, on the 
average 3.5 km [sic] (Yanushko, 1957; and others). 

Judging from the few observations made in the Altai, western 
Sayan, and central Sikhote-Alin, where winter is extremely snowy, 
one may assume that deer at that time of year are confined to even 
more restricted areas than in the European part of the USSR. In the 
event of ice crusts the animals cannot go beyond two to three hec- 
tares for a few days at a stretch. In Caucasian preserve in the mid- 
1930's it has been stated that the home range of a male in rut with a 
harem of females usually did not exceed 400 to 500 hectares (Nasi- 
movich, 1941). 

Daily activity and behavior. In hot weather in summer deer stop 
grazing by 8:00 or 9:00 a.m. (as soon as the dew dries); grazing is 
resumed around evening and extends, with intervals, into the night. 
In cloudy weather and light rain, especially when the animals are 
not frightened, grazing deer can be seen at all hours. '''^ For precisely 
the same reason, high in the mountains where it is much cooler, 
many deer graze even during daylight hours, as observed in Cauca- 
sian preserve. 

In summer relatively open and windy areas (ridge crests, open- 
ings in woods, slightly sprouted burned-over forest sections, river 
shoals, lake banks, etc.) as also dense, well-shaded thickets of tall 
grass, shrubs, etc. serve as beds and sites for cud chewing, depending 
on the time of day, air temperature, abundance of various blood- 
sucking insects, etc. In Caucasian preserve female deer quite often lie 
in such tall grass that they are completely hidden; males do not 
miake beds in such places. In large burned-over forest sections deer go 
to a lake (alpine segment of western Caucasus) or a river (the Sayans) 
and sometimes lie on snowfields when midges are prolific. To save 
152 themselves from mosquitoes and gadflies deer of Azov-Sivash pre- 
serve descend into the water of bays and lie with only their heads 
above the water (M.D. Shpigov). 

They usually do not graze during heavy rains. 



204 




Fig. 50. Summer habitat of deer in the Caucasus. Subalpine glade on Abago 
mountain. July, 1946. Caucasian preserve. Photograph by I.V. Zharkov. 



Bukhara deer graze mainly at night in summer, usually far away 
hom rivers (where mosquitoes are prolific) at the border of deserts; 
during the day they rest in forests (not avoiding saline sections); 
when there is a rise in water level they lie directly in the water 
(Flerov, 1935). When threatened by mosquitoes, deer of Dargana- 
tinsk tugais quite often enter neighboring desert sections at night. 
Night grazing in the tugais is interspersed with periods of rest 
(Klyushkin, 1949). Deer visit salt licks and waterholes mainly during 
hours of darkness, but in Caucasian preserve they are often seen in 
salt licks during the day as well. 

In winter, especially if there is a food shortage, deer spend most 
of the day feeding and, if not threatened, the activity rhythm 
depends not so much on the time of day as the weather conditions. 
During frosts, without wind, deer are generally more active than in 
mild weather; they try to protect themselves from the wind in 
ravines, more frequently in shrubs, among dense coniferous planta- 
tions and sometimes in caves (Crimea). They remain in their beds 
during intense snowfall. In winter sunny slopes with open vistas are 
a favorite place for beds in mountains. In valley sections where the 
wind blows rather continuously deer usually do not make beds; 
quiet places are invariably preferred (Kaplanov, 1948). 

The weight load on the hooves of deer is considerable; in adult 
animals it is 450 g per cm^ (Voronezh preserve; Mertts, 1957); in 



205 

Crimea, evidently, it is much more (Yanushko, 1955). Thus with an 
increase in height of snow cover deer movement becomes increas- 
ingly difficult. In order to ease their movement on snow deer utilize 
their earlier tracks and walk under dense crowns of coniferous plan- 
tations. In addition they reduce their mobility and feed intensely at a 
single place at such times. The critical height of snow cover for 
adult Altai maral is about 70 cm; the corresponding value for most 
other forms of red deer is 50 to 60 cm. In their search for items of 
food such as plant remains, acorns, beechnuts, etc. deer dig up the 
snow with their muzzle (if the snow is loose) and hooves, but only as 
long as the snow cover is less than 25 to 30 cm deep. 

Deer move quite fast and can cover 4 to 5 km in an hour. Fright- 
ened, they immediately gallop but rarely trot; the span of a jump can 
exceed 6 m. Deer are excellent swimmers and instances are known of 
their swimming through turbulent (Terek and Pyandzha) and broad 
(Lena) Rivers or, to save themselves from wolves, even swimming 
several kilometers into the sea (Abramov, 1954). 

Olfactory and auditory senses are well developed in deer; vision 
is evidently equally good but the animals are not so dependent on it. 
While feeding, the head is usually held high in the air. Instances are 
known of a sleeping deer (bedded or standing) walking off almost 
instantly on being awakened. 

In regions where winter is relatively snowy the gregarious ten- 
dency is more pronounced in the cold period of the year than at any 
other time. Not only are mixed herds formed (more common in 
winter but the number of adult males in them is almost invariably 
small), but also herds consisting exclusively of males or females with 
young (herds of the second type contain a larger number of ani- 
mals). In winter mixed herds are often formed from autumn harems 
with fawns and juveniles of ages 1.5 to 2.5 years, which had separ- 
ated in the period of rut. 

In spring, before breeding, mixed herds as well as those compris- 
ing adult females and young break up. Many small herds are then 
formed from the "large family" of an adult female (playing the role 
of a leader) and the offspring raised over several years. 

Herd size depends mainly on overall deer population, amount of 
food in the region, etc. The usual herd size is three to six animals, 
rarely more (in the western Caucasus from the end of the nineteenth 
to early twentieth century deer herds consisting of up to 70 animals 
were recorded; Dinnik, 1910). Even in Crimean preserve, which 
abounds in deer, about 60% of the entire population is contained in 
herds of 4 to 10 animals each, with 30% in smaller herds and only 
10% in larger ones (Yanushko, 1957). 



206 




Fig. 51. 



Deer track. Voronezh preserve. March, 1958. Photograph by P.F. 
Kaznevskii. 



Seasonal migrations and transgressions. Depending on the 
nature of biotopes, food availability, distribution and thickness of 
snow cover, and other factors, deer winter in the same locality where 
they lived in summer, only changing over from one type of habitat 
to another (Belovezh Forest and Voronezh preserve). Sometimes they 
undertake more extensive migrations, especially in montane regions 
(Carpathians, Crimea, the Caucasus, southern Siberian mountains, 
and Sikhote-Alin). The maximum number of deer congregate for 
154 wintering in regions where the snow cover does not exceed 40 to 60 
cm, but adult males often winter in mountains where the snow cover 
is 100 cm or even thicker, provided there is abundant twiggy food 
(western Caucasus and Altai). Wintering in such places is facilitated 



207 



by the absence of wolves, which congregate in less snowy regions 
(Derevyanko andZharkov, 1953). Since females, fawns, and juveniles 
choose wintering places where the snow cover does not exceed 40 
cm, places of mass wintering of females and adult males often do not 
coincide; the latter usually locate at higher altitudes. 

In the mountains, in addition to downward vertical migrations 
in autumn, winter migrations to wind-protected, leeward slopes 
occur in less snowy mountain belts (range of vertical migrations 
may go up to 1 ,000 to 1 ,500 m). Such slopes provide a more favorable 
situation with relation to snow than excessively snowy windward 
slopes (Altai, the Sayans, Kuznetsk Alatau and Barguzinsk ranges). 
Having gathered in such leeward sites for wintering, the animals 
often cross several parallel ranges for feeding. The distance covered 
during wanderings in some cases ranges from 50 to 140 km (Nasimo- 
vich, 1955). 

Migration to wintering sites usually occurs gradually and 
extends for a period of 1 .5 to 2.0 months and sometimes even longer. 
More often, movement commences in the period of rut or imme- 
diately after it (rarely before), i.e., from October, and ends in 
November (more rarely in December- January). The first to reach the 




154 Fig. 52. Two-year-old female maral. Mouth of Ad-Kicha River near Lake Telets. 
Altai preserve. End of February, 1951. Photograh by F.D. Shaposhnikov. 



208 

wintering sites are females with young. Heavy snowfalls accompan- 
ied by cooling and seasonal changes in the landscape serve most 
often as a signal for the commencement of mass migrations. Under 
these conditions, when the snow thickness exceeds 20 cm, it becomes 
difficult for deer to dig up plant food from under the snow. 

In the Caucasus and Sikhote-Alin in the second half of winter, 
when snow is heavy, secondary migrations of deer occur down the 
155 slopes into less snowy sections (Nasimovich, 1939; Kaplanov, 1948). 
In the event of formation of ice crusts deer congregate along well- 
shaded slopes in fir, spruce, and cedar forests. 

The return to summer habitats commences in the period of 
intense snow thawing and is over in May or early June. The migra- 
tory paths of deer in many regions are fixed and coincide sometimes 
with those of elk and roe deer. 

Summer dispersal and migration of deer depend greatly on the 
availability of food, shelters in the locality, and absence of blood- 
sucking insects. For example, in the Sikhote-Alin mountains with 
the appearance of midges (June) many Manchurian wapiti, espe- 
cially the males, go high into the mountains and congregate there in 
deciduous, burned-over forest zones near glades, where blood- 
sucking flies are few because of the openness of the expanse and 
stronger winds. In the latter half of summer, with the onset of 
intense heat and the mass appearance of gadflies, these deer desert 
the open mountains and congregate in cool, wet, and shaded places 
(bottomland deciduous forests, river drainages, alder thickets, and 
boreal forests) where they suffer less from gadflies and heat 
(Kaplanov, 1948). 

The small population of Bukhara deer performs only local, 
migrations. In the Vakhsh basin the manifestation of such migra- 
tions is extremely poor; in spring and summer when the water level 
rises deer only desert to drier islands, continuing to wander along 
submerged areas (Flerov, 1935). Migrations caused by floods are seen 
in Darganatinsk tugais. In other cases migrations of deer are mostly 
related to agricultural and other activities of man. On being con- 
stantly exposed to threats deer often migrate from one set of tugais to 
another or even move off into the desert for some time (Klyushkin, 
1949). In the Syr-Darya basin Bukhara deer have now been almost 
exterminated. Here they formerly spent much of the summer in 
saxaul thickets. In August, due to scarcity of water in Kyzylkum, 
deer came close to the rivers and were confined to the tugai zone; in 
November, at the commencement of snowfall, they again went into 
the desert (Bobrinskii, 1933). 

Instances of reappearance of deer in regions from where they had 



209 

disappeared long ago are a frequent phenomenon in many parts of 
their geographic range. Such instances have been recorded time and 
again in the northern and eastern Pri-Caspian (Bazhanov, 1945), 
northern Caucasus, '°* and other places. In Bashkir preserve, where 
maral had acclimatized, their appearance in new habitats usually 
occurred in the latter half of winter and was associated with food 
scarcity at that time of year, leading to wanderings. In snowy winters 
(1941-1942, 1942-1943, 1947-1948, and 1949-1950) intrusions into 
new habitats have assumed mass dimensions; some of the animals 
settled down in new places and did not return to old habitats in 
spring. The range thereby enlarged gradually and maral dispersed 
mainly along river watersheds covered with sparse pine forests and 
later even occupied bottomland areas (P.F. Kaznevskii). 

In the winter of 1953 to 1954, during snowstorms on Biryuchii 
Island (Azov-Sivash preserve), some deer reached Arabatsk Strelka 
(Crimea), covering a distance of up to 30 km on ice (Ishunin, 1956). 

Reproduction. Deer come into rut in autumn. It is manifested in 
some males externally by bellowing [bugling], initially not so 
loudly and for only a very short period, by the end of August or early 
September (in some cases from August 20 to 25). '°^ During the 
second half of September bellowing intensifies both in number of 
participants and loudness and can be heard over a distance of several 
kilometers. In most cases mass bellowing occurs in the second half 
of September but sometimes in October also. In the latter half of 
October bellowing ceases, but some stray males in some years can be 
heard in November and, in exceptional cases, even in December- 
January. 

During the period of maximum bellowing deer commence with 
broken and hoarse sounds resembling heavy breathing ("ahh-ahh- 
ahh") followed by a low, powerful, and prolonged mooing ("ee-ah- 
ooo-ooo-oooo") in which the last syllable in males of up to 2.5 years 
of age resembles a howl. Among Voronezh, Crimean, Caucasian, 
and Altai deer the style of bellowing is generally similar but the 
pitch in some cases higher than in others. Manchurian wapiti and 
Bukhara deer are characterized by extremely low and hoarse 
bellowing. 

The bellowing period generally varies from one to two months; 
there are, however, extreme variations in different years in duration, 

'"^For example, in 1956 a small herd of deer was sighted in an "island" of forest in 
the plains near Crimea station in northwest Krasnodar region, 50 to 60 km 
away from the nearest mountain (A.V. Vinogradov). 

'"'in an extremely rare case the weak bellowing of a lone deer was heard even on 
August 15 and 16 (Sikhote-Alin and Crimea). -^ 



210 

harmony, and intensity of bellowing. Weather, feeding, and other 
seasonal conditions exert great influence on the nature of bellowing 
and rut. Following the extremely snowy winter of 1944 to 1945 Man- 
churian wapiti in Sudzukhin preserve bellowed very little (G.F. 
Bromlei). In 1908 deer of western Caucasus suffered from foot-and- 
mouth disease and did not bellow at all that autumn (Dinnik, 1909). 
Even in normal years part of the adult male deer population (usu- 
ally not less than 20 to 30%; Zharkov, 1952; and others) will not 
bellow. If the sex ratio in a population tends to approach 1:1, com- 
petition for females intensifies and so, too, does bellowing. In popu- 
lations with a significant predominance of females bellowing is 
usually less intense (Mertts, 1954; and others). 

Commencement of bellowing and the subsequent course of rut 
are largely predetermined by phenological conditions: coloration of 
the landscape, onset of frosts, and so on (Sablina, 1955; and others). 
A perceptible drop in air temperature around the time bellowing 
should commence activates the males. If cooling is accompanied by 
inclement weather, prolonged rains, etc. after deer have been bellow- 
ing for some time, its intensity drops or it ceases altogether. If the 
autumn season is rainy throughout, bellowing is dull, unenthusias- 
tic, and extended for a longer than usual period (Nasimovich, 1936 
and 1941; Kaplanov, 1948; Yanushko, 1947; and others). Deer bellow 
mostly in the evening and morning but at the peak of rut through- 
out most of the night, especially on moonlit nights (Nasimovich, 
1941). 

In Crimea mass bellowing of deer usually occurs from September 
21 to 26 (Yanushko, 1957). In Caucasian preserve mass bellowing 
has been recorded in various years from September 20 through 
October 5 (Nasimovich, 1941). A sizable number of deer commence 
bellowing earlier, from September 15, in the Laba basin where the 
mountains are high and the climate more continental. In the Beloe 
basin full-throated bellowing usually commences a few days later. 
In the Golovinka basin, on the southern slope of the Great Caucasus 
range, it commences from September 25 only (Zharkov, 1952). In 
Zakatal'sk and Lagoda preserves, where the climate is much milder 
than in Caucasian preserve, mass bellowing of deer usually occurs at 
the fag end of September and in the first ten days of October (I.E. 
Popkova and G.P. Enukidze). 

In Altai preserve peak bellowing among maral has been recorded 
from September 16 to 17 through 22 to 24, lasting usually for five to 

""According to Kotovshchikova (1936) Crimean deer located in lower montane 
altitudes begin bellowing earlier. 



211 

seven days (Dmitriev, 1938). In the middle part of Sikhote-Alin, mass 
bellowing of deer is most often heard from September 20 through 25, 
and in southern Primor'e region from September 25 through 
October 1 (Abramov, 1954). 

References to periods of rut among Bukhara deer are contradic- 
tory. According to Flerov (1935 and 1952), Klyushkin (1949), and 
other researchers bellowing commences mid-September and some- 
times even later (in Darganatinsk tugais often only from early 
October) and ceases in the latter half of October or even early 
November. Deer commenced bellowing in Tigrovaya Balka preserve 
from 1946 to 1950 from August 30 to September 6 and ended Sep- 
tember 26 to October 5; mating was observed up to October 10 (Cher- 
nyshev, 1958). In Moscow Zoological Garden bellowing of Bukhara 
deer peaked in the first or second half of October — a delay of two or 
three weeks compared with maral (Tsalkin, 1944). 
157 At the very commencement of bellowing many males are without 
females and remain alone, occupying a definite home range, some- 
times the same one for two or more consecutive years (Caucasian 
preserve).'" Sexually aroused males strike small trees with their 
horns, peel off the bark, and break branches and tops. By stamping 
with its hooves each male creates two or three "tochkami" (so-called 
by hunters)* of bald turf within its territory, sometimes saturating 
them with urine. Wallowing in mud iд their bathing regions is also 
common among males in rut. In the second half of September many 
males (at the peak of rut not less than 50% or more) move with two or 
three females or sometimes more. In the mountains of the northwest 
Caucasus males with up to 10 females each have been sighted and in 
one instance in Crimean preserve a male with 19 females (including 
juveniles). In maral farms males live with 4 to 15 or even up to 20 
females (Zalesskii, 1930). 

Males are capable of mating from the end of August (or slightly 
earlier) up to November and in some cases even later. In Crimean 
preserve individual males with traces of spermatogenesis have been 
caught in almost all months of the year. Mating is maximal from 
August to November, especially in September and October, after 
which it gradually declines (in April and May the tests are relatively 



The manner of bellowing of some males is so specific that they can easily be 
identified even in different years. 

"^Most juveniles (up to 1 .5 years) live separately, away from adults, in the period of 
rut. 

*Literally, "points" — Sci. Ed. 



212 

inactive); sexual activity resumes in July or even earlier. The forma- 
tion and development of follicles in the ovaries of adult females 
occurs throughout the year but Graafian follicles attain full matur- 
ity and ovulation occurs only in the period of estrus (Yanushko, 
1957). 

Mating usually takes place not earlier than mid-September, more 
often slightly later. In the middle part of Sikhote-Alin most Man- 
churian wapiti mate from mid-September to mid-October and in 
southern Primor'e region from September 25 through October 20 
(Abramov, 1954). In Moscow Zoological Garden mating has been 
observed in the period of maximum bellowing and later, for a period 
of a month or more; most maral mate here from the end of Sep- 
tember to early October, while most Bukhara deer mate later, in 
mid-October (Tsalkin, 1944). Mating is rapid and takes place while 
the animals are moving; a single female is usually covered several 
times. 

There is no doubt that bellowing facilitates the location of males 
by females and at the same time exerts a stimulating influence on 
female estrus. During bellowing deer remain in places where their 
voice will carry long distances. The intensity of bellowing is inver- 
sely proportionate to harem size; males with large harems bellow 
less than males which, for some reason, are haremless. With the 
approach of a stronger competitor a male with a harem usually falls 
silent or even attempts to run away with his females (Kaplanov, 
1948; Mertts, 1953; Abramov, 1954; and others). Deer judge the 
strength of a competitor not only from his external appearance but 
also from the nature of his bellow. In young males the bellow is very 
high pitched, while in virile animals it is very low and hoarse. There 
is no doubt, therefore, that weak bellowing is associated with fright 
and cannot be regarded as a "challenge to arms". Young males up to 
the age of 2.5 years hearing the call of a large male try to hide. In 
most regions fights over a female are not infrequent between adult 
males and may be violent at times, ending in the death of one or 
both rivals. Instances are known of antlers being broken or so 
entangled during a fight that the combatants could not separate" 
themselves. Very often the approach of a rival is restricted to threats; 
the deer dash their antlers against trees but do not attack each other. 

Some herds with harems are large, probably formed by the 
merger of two or more harems, and thus contain not one but two, 
three, or even more males. This phenomenon is relatively widely 
158 prevalent in Crimean preserve where density of deer is very high and 
individual harems less isolated from each other than in other 
regions. Thus in Crimean preserve (1950 to 1954) 77% of all the 



213 




158 Fig. 53. Deer killed in a duel during the period of rut. Voronezh 

preserve. September 25, 1950. Photograph by P. A. Mertts. 

harems sighted included one male, 18%, two males, and 5% three, 
four, or more males (Yanushko, 1957). Evidently larger herds with 
more than one male are not very stable and readily split into smaller 
ones with a single male in each. 

Males bellowing actively (loners or those with small harems; 
Mertts, 1954; and others) eat little"^ and lose up to 30 to 40 kg or 20 
to 25% of their summer weight during rut (Heptner and Tsalkin, 
1947; Flerov, 1952). Judging from the results of mass shoots male 
deer in Crimean preserve lost during rut an average of 16% summer 
weight, but later recouped rapidly (Yanushko, 1957). At places 
where the climate is more severe, winter sets in early, and nourishing 
foods (for example, acorns) are not available every year, some males 
remain emaciated until winter. If in addition the winter is snowy 
and severe, mortality among adult males will be significantly higher 



In such deer the stomach contents might not exceed 0.2 liters (Mertts, 1954). 



214 

than among females or even juveniles (Merits, 1953 and 1957). 

Among Crimean deer spermatogenesis commences from the age 
of 17 to 19 months, when males cannot yet be regarded as adults 
either in weight (75 to 123 kg), height at withers (96 to 118 cm), or 
structure of teeth and antlers. These deer do not participate in breed- 
ing and ordinarily do not bellow. Most males commence bellowing 
in their third year but even then do not usually participate in breed- 
ing because of competition from more powerful adult animals 
(Yanushko, 1957 and 1958). These data are based on a large amount 
of factual information and are evidently true for species of red deer 
as a whole. In maral nurseries instances are known of three-year-old 
males being mated with females who later fawned; usually males 
mate successfully only at the age of five or six years and rarely four 
(Zalesskii, 1930). 

Among Crimean deer a few individual females come into heat in 
their second year when growth has not yet ceased;^''* most females, 
however, bear their first offspring only by the end of the third year; 
about 70% become pregnant in their third year during the period 
October through May (Yanushko, 1957 and 1958). In Latvia about 
50% of all young females are usually impregnated in the third year, 
and by the end of that year bear their first fawn (Kalnin'sh, 1950). 
Similar data on periods of sexual maturity have been given for deer 
in Belovezh Forest (Sablina, 1955) and several other parts of the 
159 range. According to the data of mass shoots of deer in Germany the 
number of pregnant females in their second year (total of 1,739 
animals) in different regions varied from to 63% or an average of 
49% (Kroning and Vorreyer, 1957). Thus in the western sections of 
the range a significant percentage of females participate in mating 
and reproduce even in the second year. Possibly this should be consid- 
ered the result of selection, fattening, and other measures adopted 
in game farms. In maral nurseries females become pregnant in the 
fourth or third year and rarely in the second. 

Some adult females do not fawn every year (due to late fawning, 
sickness, age, etc.). The proportion of such females in the total 
number of mature females may reach 25% (Crimean preserve) or even 
44% (Voronezh preserve) (Yanushko, 1957 and 1958). In populations 
of west European deer and American wapiti 20 to 25% are usually 
barren (Trippensee, 1948). Instances are known when females in 
maral farms fawned for four to six consecutive years; even then 
instances of barrenness were not uncommon (Zalesskii, 1930). 

Young females attract attention later than adults and hence some are impregnated 
late in autumn or even in winter (Merits, 1954). 



215 

Gestation usually lasts 34 or 35 weeks but is probably somewhat 
longer in isolated cases. Delayed implantation is absent and the 
embryo is discernible even at the end of November (Caucasian pre- 
serve). Among most subspecies of red deer the majority of females 
fawn in the latter half of May (often only from the end of that 
month) or in the first half of June (more often up to June 10), but 
some individuals which have been impregnated much earlier, or on 
the contrary later, fawn even in the first half of May or at the end of 
June to early July.^'^ Yet instances are known of isolated females 
attracting attention at an unusual time, mating, and bearing off- 
spring very early, even by the end of March (Crimean preserve, 1947), 
or extremely late, in the latter half of November (Birobidzhan, 1925; 
Voronezh preserve, 1949). Significant deviations from normal fawn- 
ing periods are frequent among maral. Bukhara deer fawn from the 
end of April through May inclusive (Flerov, 1935; Chernyshev, 1958; 
V.I. Chernyshev). In Moscow Zoological Garden fawning mainly 
takes place in June (Tsalkin, 1944).''^ 

Females fawn in secluded spots, often in fern thickets or other 
vegetation along banks of rivers and springs. In an overwhelming 
majority of cases births are single. In Crimean preserve, of the 149 
pregnant animals studied only one female with two embryos was 
found (Yanushko, 1957). In Azov-Sivash preserve from 1948 to 1953 
twins were found only once in 71 births (G.I. Ishunin). In Altai 
maral nurseries roughly two sets of twins are born in every 100 
recorded births (Zalesskii, 1930). 

Growth, development, and molt. At birth a fawn* is weak and 
helpless. Attempts to stand are made in about half an hour after 
birth but usually are not successful in the first few hours; the animal 
has hardly stood before it falls down. At the age of three hours fawns 
of Bukhara deer can stand for some time and commence suckling 
(Tsalkin, 1944). For the first two or three days fawns lie down nearly 
all the time and stand up only to suckle. The mother moves a dis- 
tance of 100 to 200 m to graze and feeds her young several times a 
day; at other times she usually stands apart to render detection of the 
fawn difficult by predators. From the fifth to the seventh day fawns 
begin to trail their mothers but are not stable on their legs even 



In Azov-Sivash preserve over 10% females fawn in July, possibly because the deer 
population there is hybrid (G.I. Ishunin). 

Of 27 calvings among Bukhara deer in Moscow Zoological Garden, none was 
earlier than June 16 and some (37%) occurred in July (Tsalkin, 1944). 
♦Usually called "calf" in North America — Sci. Ed. 



216 

though they can run. At that age fawns suckle four to six times a 
day. Two-week-old fawns run and jump well; by July they are 
almost equal to adults in running. They begin grazing at the age of 
one month and soon after begin to chew the cud (Dobrzhanskii, 
1928; Tsalkin, 1944; and others). 

The young continue to suckle at least until early winter (there is 
often a gap in the period of estrus) and in some cases much longer. 
In maral farms pregnant females drive away the young from 
December onwards, while barren females allow the young to suckle 
until the next fawning and, should the fawn die, even longer. 
160 Instances are known of females suckling newborn and one-year-old 
fawns simultaneously (Zalesskii, 1930; Tsalkin, 1944). Lactation 
before the birth of offspring is a frequent phenomenon among Lat- 
vian deer (Kalnin'sh, 1950). A few hundred deer were caught in 
Crimean preserve from 1951 to 1953. Among the females caught 
from July to September those lactating represented 74%, from 
October to December— 69%, from January to February— 41%, and 
from March to May— 32% (Yanushko, 1958). Juveniles remain with 
the mother up to the age of 1 .5 to 2.5 years; males usually separate at 
the age of 1 .0 to 1 .5 years; some young females, however, even after 
their first fawning rejoin their mothers. 

Judging from the very few weighings of newborn deer from dif- 
ferent parts of the range, their weight varies from 8 to 11 kg. The 
young grow very rapidly and put on weight for about six months; 
thereafter growth, especially in females, slows down and at the same 
time sexual dimorphism in size intensifies (P.A. Yanushko). In 
maral nurseries three-month-old male maral weigh 46 to 67 kg (M 
55); one-year-olds weigh 120 to 138 kg (M 130.6); two-year-olds 189 
kg; three-year-olds 231 kg; four-year-olds 248 kg; five-year-olds 262 
kg; six-year-olds 280 kg; and seven-year-olds and older 297 kg. By the 
sixth or seventh year weight increase ceases or becomes imperceptible. 

Manchurian wapiti, as well as other subspecies of red deer, are 
inferior to maral in size; male adults weigh 170 to 250 kg and females 
140 to 180 kg (Kaplanov, 1948). Judging from numerous weighings 
of Crimean deer, adult males in rare instances reach 269 kg and 
females 166 kg. On the average, however, the weight difference 
between adult males and females is about 50 kg (P.A. Yanushko). 

Toward the end of the first year small bony prominences appear 
on the frontal bones of males. These prominences gradually grow 
into stumps which later serve as bases for antlers. The antlers them- 

"'According to Tsalkin (1944) sexual dimorphism in the size of maral is well 
manifested even by the third month. 



217 

selves begin to grow from 12 to 14 rnonths; by the autumn of the 
second year growth ceases, ossification sets in, and the velvet is shed. 
These first antlers in an overwhelming majority of cases are devoid 
of tines and are variously called spikes or awls; these names are also 
applied to young with such antlers. The first antlers are usually shed 
in April, after which antlers with three or four tines begin to grow. 
In subsequent years the dimensions of antlers and their weight and 
number of tines increase (the tines do not increase in number until 
the animals are five or six years old or in the case of Bukhara deer 
four or five years old). Deer in the age group of 6 to 8 to 12 to 14 years 
have the best developed and heaviest antlers (in rare cases up to 16 
and even 18 years of age), after which deterioration sets in with a 
reduction in size, number of tines, and weight. The weight of nor- 
mally developed antlers of maral reaches 7 to 10 kg and in rare cases 
18 to 20, or even 24 kg. Among Caucasian deer antlers weighing 7 to 
8 kg are not a rarity, sometimes reach 10 to 11.5 kg, and exception- 
ally 14 to 18 kg. Adult Manchurian wapiti in Moscow Zoological 
Garden have antlers weighing 3.5 to 7.0 kg and Bukhara deer antlers 
weighing 3.5 to 5.5 kg (Turkin and Satunin, 1902; Dinnik, 1910; 
Tsalkin, 1945; Heptner and Tsalkin, 1947; Astanin, 1949; and 
others). 

In most regions the majority of deer shed their antlers from the 
second half of March to early April. In the Baltic, Carpathians, 
Crimea, Azov-Sivash preserve, and the Caucasus, at least in some 
years, many deer shed their antlers even at the end of February (in 
Crimea some loners which have shed their antlers are sometimes 
observed from the end of January to early February; P.A. Yanushko). 
In Belovezh Forest shedding usually occurs only from the tenth of 
March. Maral in Bashkir and Altai preserves and Manchurian 
wapiti in the Far East shed their antlers only in the second half of 
March, often at the end of the month. References to periods of antler 
shedding among Bukhara deer are contradictory. Some stray deer, 
usually very young, shed their antlers later than virile stags and are 
161 seen with old antlers until the end of April or early May, and in rare 
cases until the end of May or early June. In Moscow Zoological 
Garden delay in shedding of antlers by deer 2 to 4 years old com- 
pared to deer 5 to 13 years old, reaches almost 1.5 months. Shedding 
of antlers is also delayed in old deer (15 to 18 years), but not to the 
same extent as in young ones (Tsalkin, 1945). In years of severe 
winters antler shedding occurs later than usual (Bashkir preserve; 
Kirikov, 1952). 

New antlers commence growing within a few days of old antler 
sloughing. New antlers are sheathed in a tender hairy skin, soft, and 



218 

initially very sensitive not only to injury but also to mosquito bites 
(Przewalski, 1870). They are termed "velvet" antlers. In most deer 
antlers are tully formed by the end of June or the first half of July. 
After this they become ossified (July end to the first half of August) 
and the skin ("moss") is shed. This process is completed first in 
Crimean and Caucasian deer and later in maral and Manchurian 
wapiti (Tsalkin, 1945). For the first few days after the velvet is 
sloughed the antlers are light-colored but darken thereafter. Referen- 
ces to periods of antler formation among Bukhara deer are contra- 
dictory. The ossification of antlers according to various researchers 
ceases by the end of July to mid-Augusr and sometimes only at the 
end of September in the young. A 2.5-year-old male was caught in 
the Pyandzha in mid-August in which the antlers were still not 
completely formed and unossified (Flerov, 1935 and 1952). 

In Moscow Zoological Garden the duration of growth of antlers 
among adult deer of various subspecies is similar regardless of antler 
size or stniciure. The process of antler formation requires 102 to 1 19 
days, after which ossification continues for 30 to 40 days. The total 
duration of antler formation therefore requires 137 to 153 days. Skin 
is sloughed from antlers in one to three weeks. Antlers ossify three or 
four weeks before the commencement of rut (Tsalkin, 1945). 

On the sides and back of young deer several rows of light-colored 
spots are visible, which gradually dull in the latter half of summer 
and disappear by autumn, after molt. Shedding of the juvenile coat 
in the first winter, essentially similar to the corresponding process in 
adults, terminates among maral of Moscow Zoological Garden in 
October, and among Bukhara deer in November (Tsalkin, 1944). In 
rare cases females continue to bear faint traces of spots even in the 
second year (Caucasian preserve). By February-March the hair has 
become intensely worn in adult deer, the beard brittle, and the fur 
lusterless and light-colored; some hair in the beard begins to fall in 
March. In much later stages of molt the underfur is usually shed, 
falling out in large tufts which cling to bushes when deer brush 
against them."^ Deer molt most intensely from May to the first half 
of June, i.e., when relatively warmer weather sets in. The limbs and 
head are the first to be freed of winter hair, followed by the abdomen, 
thighs, sides, and back; the last to molt are the speculum, pelvis, and 
part of the neck (Tsalkin, 1946; and others). The summer beard 
begins to grow perceptibly from the end of April or in May. 

In the southern parts of the range deer molt commences from the 

Among juveniles shedding of underfur is discontinuous and it does not fall out in 
large tufts (Tsalkin, 1946). 



219 

latter half of March (Azov-Sivash preserve, Crimea, and the Cauca- 
sus), but becomes perceptible usually only in April in the northern 
regions, which are climatically more severe (Mordov preserve and 
some regions of Siberia). Molt of Manchurian wapiti in the Far East 
(Kaplanov, 1948) and Bukhara deer in Central Asia commences in 
April (Flerov, 1935)."^ The commencement of molt and its course 
depend largely on the winter regime and the physical state of the 
animal. In Crimean preserve molt after a warm winter is apparent 
by the end of March (sometimes even mid-March), but after a very 
162 cold winter evident only in April (in some years only from mid- 
April; P. A. Yanushko). The first completely molted deer in Crimea 
are observable in early May; nearly aJl deer sport a summer coat by 
the end of May, but in some years not until the first 10 days of June. 
In the more northern sections of the range and also where the moim- 
tains are significantly higher and the climate colder (Caucasus, 
Altai, and Sayan ranges), deer with large patches of winter hair can 
be seen even at the end of June or early July. In Moscow Zoological 
Garden, Caucasian and Bukhara deer molt by the end of June and 
maral and Manchurian wapiti by mid-July. The individual dura- 
tion of molt extends over 2.5 to 3.5 months, being longest among 
maral (Tsalkin, 1946). The first to molt are adult males and barren 
females, followed by three-year-olds and females with fawns. The 
process of molt is far shorter in duration among juveniles than 
among adults. 

The winter growth of beard commences at the end of July to 
early August but is only weakly discernible. By August the summer 
beard is partly shed and the growth of winter beard quite visible. 
The bright rusty summer coloration gradually changes to a rusty- 
gray and, as winter approaches, to grayish-brown. In September the 
summer hair, except on the head and limbs, is shed completely and 
the underfur commences to grow from the latter half of August, 
lengthening perceptibly. In the period of breeding deer sport transi- 
tional coats; the winter coat forms only in the latter half of October 
or early November but the lengthening of hair usually continues up 
to early December. 

Among highly emaciated males in the postrut period beard 
growth and underfur formation are greatly delayed; in the event of 
early and intense cold weather this delay notably weakens such ani- 
mals since considerable loss of body heat occurs (Mertts, 1953). 

The initial stages of molt in nature are difficult to perceive and most probably the 
reference to such a late commencement as April is erroneous. In Moscow Zoological 
Garden molt of maral, Manchurian wapiti, and Caucasian and Bukhara deer 
commences almost simultaneously in mid-March (Tsalkin, 1946). 



220 

Autumn molt generally occurs in a reverse sequence compared to 
spring molt. Initially the upper body parts, speculum, and more 
rarely sides, and later the underbody parts are freed of summer hair. 
The head and lower parts of limbs may not molt. Autumn molt is 
completed in a much shorter period than spring molt. Among adult 
maral in Moscow Zoological Garden molt lasts for 21 to 30 days and 
in juveniles for 15 to 23 days (Tsalkin, 1946). 

In the colder parts of the range autumn molt terminates earlier 
than in warmer regions, by as much as two or three weeks. In Cri- 
mea the first completely molted deer are seen from mid-October, but 
animals with incomplete molting are evident right up to the middle 
of the following month (Yanushko, 1957). 

In captivity deer may live for 20 years and in rare cases even 25 to 
27 (Kirilov, 1902; and others). '^° In German game farms some deer 
live to the age of 25 to 30 years (Severtsov, 1940 and 1941; and others). 
Cases of deer in maral nurseries yielding up to 23 sets of antlers 
during their lifetime have been recorded (Cherkasov, 1884). In pre- 
serves, left to themselves, an overwhelming majority die at 12 to 14 
years of age. Very old animals with totally worn out teeth are rarely 
encountered (Kaplanov, 1948; and others) and doomed to death in 
winter from hunger. The average life span of female deer in nature is 
unquestionably longer than that of males. 

The last milk tooth in deer is invariably shed in their third year. 
Age-related changes in dentition, by which the approximate age of 
deer can be judged, have been detailed by Raesfeld (191 1) and more 
recently by Beme (1957) for Crimean deer. 

Enemies^ diseases, parasites, mortality, competitors and popula- 
tion dynamics. Among the various causes of deer mortality over a 
vast stretch of their range, the most important are predators and 
severe snowy winters. Some diseases are equally important but their 
role in the population dynamics of deer has yet to be investigated. 
163 In the region of the Voronezh preserve between 1933 and 1952 the 
number of recorded deer mortalities was 222 (excluding those killed 
by poachers). Of these, the cause of death was established for 208;'"' 
54.3% died from emaciation during severe snowy winters, '^^ 31.2% 
were killed by wolves, 13% died from fatal wounds inflicted in duels 



'^"Kalnin'sh (1950) erroneously states that captive deer live up to 60 to 70 years. 

" Among the unascertained causes of death, disease was undoubtedly an 
important factor. 

' This category is identified easily, while that of destruction by predators often 
cannot be established; hence these figures should be taken as purely approximate. 



221 

during the period of rut, and 1.5% were drowned in rivers in spring. 
Among deaths caused by wolves (43 adult females, 20 female fawns, 
and only 2 adult males) over 90% occurred in winter months. Deer 
remains were found in winter in the stomachs and feces of wolves in 
21.2% of all the samples analyzed; the corresponding figure for sum- 
mer-autumn months was 5.3% (Mertts, 1953 and 1957). 

In Belovezh Forest over 1,100 samples of the food of wolves were 
analyzed between 1946 and 1950; deer remains were detected in 7.2% 
(in maximum quantities in autumn and winter). In some years of 
severe snowy winters (for example, the winter of 1949 to 1950), the 
number of deaths of deer caused by wolves almost doubled (Gavrin 
and Donaurov, 1954). In Caucasian preserve no less than 60% of 
juvenile deer were once killed by wolves (Teplov, 1938). 

In Altai preserve of the 107 deaths of deer registered between 1940 
and 1951, no less than 63% were ascribed to the activity of various 
predators. As a matter of fact this number should probably be higher 
since an additional 9% mortality was attributed to deer falling from 
rocks in summer, often as a result of pursuit by predators. Of the 67 
deer killed by predators, wolves were responsible for 45 and lynxes, 
wolverines, and bears for the rest (Dul'keit, 1956). 

Adult deer are chased by packs of wolves or at least a pair; a lone 
wolf finds it difficult to cope with a deer, especially a stag. To 
protect themselves from attack deer lift the anterior part of the trunk 
and strike with the front hooves by extending them forward; males 
also make use of antlers in combat. Deer in montane regions attempt 
to rescue themselves from wolf attack by sheltering in rocks, in rapid 
river currents at relatively shallow spots, and when on the coast by 
swimming into the sea. 

Compared with wolves the role of all other predators in destruc- 
tion of deer is negligible. Apart from the wolf, deer are attacked most 
often by lynx, but a large number of such attacks are confined 
mainly to montane regions. Data on feeding by lynx in Caucasian 
preserve revealed deer remains in 8.3%^^^ and in Belovezh Forest in 
1.0% (Yurgenson, 1955). In Altai preserve of 56 instances of quarries 
chased by lynx 8 were maral. Family groups or pairs of lynx attack 
maral on snow; in one case they wounded an adult on snow crust 
(Dul'keit, 1953). In the eastern Altai and western Sayan, where win- 
ter is extremely snowy, maral attacks by wolverine are not rare (Ko- 
zhanchikov and Kozhanchikov, 1924; Dul'keit, 1953). Leopard and 
tiger are so few in number now that the damage inflicted by them on 

'^'Attacks on wild ungulates by lynx have increased in recent years in Caucasian 
preserve (Kotov, 1958). 



222 

deer IS quite insignificant; formerly deer attacks by them were a 
common phenomenon. In the Sikhote-AHn mountains among the 
quarries of tiger Manchurian wapiti used to occupy second place to 
wild boar (Kaplanov, 1948). Between 1946 and 1950 in Tigrovaya 
Balka preserve (Tadzhikistan) 19 Bukhara deer were killed by tigers; 
recently this deer has taken second place to wild boar in the food of 
local tigers (Sludskii, 1953). Brown bear/^" yellow-throated marten, 
striped hyena, jungle cat, wildcat, jackal, and red wolf attack deer 
164 sometimes, mainly the young, but on the whole the damage inflicted 
by them is negligible. In Caucasian preserve the death of a juvenile 
female deer was recorded as due to attack by two large predatory 
birds, evidently golden eagles (Nasimovich, 1936). 

Deer suffer from foot-and-mouth disease, Siberian ulcers, cattle 
plague, infective abortion, necrobacillosis, necrotic stomatitis (agent 
Spherophorus necrophorus), pyroplasmosis, pasteurellosis, tuber- 
culosis, leptospirosis (recorded in Slovakia), paratyphoid, and other 
diseases. Diseases in most cases were established not only in zoologi- 
cal gardens and maral nurseries, but also and usually under natural 
conditions. Epizootic foot-and-mouth disease has been detected time 
and again among deer of the northwest Caucasus (1908, 1911, and 
1925), causing suffering and death among many animals. In those 
same years innumerable sick deer fell prey to wolves, being unable 
to run away quickly. Additionally, a high percentage of the survi- 
vors did not undergo rut. Epizootic Siberian ulcers have time and 
again caused heavy losses among maral and Manchurian wapiti in 
the Altai and eastern Siberia, both in maral nurseries and among 
wild animals (Turkin and Satunin, 1902; Novikov, 1937; and 
others). In the pre-Revolutionary period, Siberian ulcers wiped out 
maral farms in Trans-Bailkal (Lezhnin, 1922). This same disease has 
been detected in Belovezh Forest. In 1886 a large number of wild and 
domesticated Manchurian wapiti died of cattle plague in Trans- 
Baikal (Kirilov, 1902). In 1911 epizootic pasteurellosis raged among 
deer of Belovezh Forest causing multiple deaths (Vrublerskii, 1912). 
Many instances of fatal diseases among deer remain undiagnosed 
(Novikov, 1937; Kolushev, 1955; and others). In western Europe 
rabies has been observed time and again among deer (Kovach, 1957; 
and others). 

In Crimean preserve two deer mortalities have been established 
as due to theileriasis caused by the blood parasite Theilena (Rukh- 



'^"in western Sayan, after wolverine, bear is considered the chief enemy of maral 
(Kozhanchikovand Kozhanchikov, 1924; Belousov, 1934). In this region wolves are few 
or altogether absent because of heavy snow. 



223 

lyadev, 1948). The same parasite has also been detected in the blood 
of Bukhara deer (Chernyshev, 1958). The protozoan Eimeria, caus- 
ing coccidiosis, has also been found in deer. 

On the whole, among deer of the Cervus elaphus group 65 spe- 
cies of helminths, including not less than 48 within the Soviet 
Union, have been identified (Belyaeva, 1958). In Crimean sanctuary 
alone 41 species of helminths have been reported for deer (Rukhlya- 
dev, 1948; Kadenatsin, 1958). Among the most pathogenic and 
extensively prevalent helminths are Dictyocaulus and members of 
protostrongylids, causing grave pulmonary disorders (Boev, 1957). 
Onchocerca bovis infects leg joints, leading to lameness; this disease 
is not uncommon among deer of Caucasian preserve (D.P. Rukhlya- 
dev). Infection of deer by liver flukes has been recorded in western 
Europe (Linke, 1957) as well as in Belovezh Forest. However, a lesser 
number of deer in the Forest have suffered from fascioliasis than 
European bison (Vrublevskii, 1927). 

Parasitism of deer by the itch mite {Sarcoptes scabei) causes sca- 
bies (Kovach, 1957); in the Soviet Union this disease has been stu- 
died little. Deer are also parasitized by gadflies — nasopharyngeal 
gadfly (P haryngotnia picta and Cephenomyia auribarbis) (Grunin, 
1957) and skin gadfly (Hypoderma). In May-June the larvae of P. 
picta were found in all 40 Manchurian wapiti examined in Sikhote- 
Alin preserve prior to 1950 (Kaplanov, 1948). Deer also suffer inten- 
sely from various blood-sucking arthropods, gadflies, mosquitoes, 
midges, Lipoptena cenn, mites {Ixodes, Haemaphysalis, Dermacen- 
tor, etc.), and other insects. A few hundred mites {Haemaphysalis 
concinna and Ixodes ricinus) have been found in some deer of Cri- 
mean preserve from April through July (Mel'nikova, 1953). Hair lice 
{Trichodectes), deer lice {Pediculans cenn), and other lice also para- 
sitize deer (Linke, 1957). 

In regions with relatively severe and snowy winters the loss of 
deer is more in years when the snow cover significantly exceeds the 
average height and persists for a prolonged period, or in the second 
half of winter when an ice crust forms on its surface. In deep snow 
(65 to 70 cm or more) young deer break the crust with their hooves 
165 and bog down in it without solid support under their legs. Even 
when the crust is so firm as to support them, adult deer often slip 
when running to save themselves from wolves, fall down, or cannot 
negotiate sharp turns (Mertts, 1957). Getting around becomes more 
complicated when the population of forest ungulates is high, result- 
ing in a shortage of tree and shrub food. In this respect the history of 
deer of Voronezh preserve, is extremely interesting: initially, when 
the deer population was low, mostly juveniles suffered in snowy and 



224 

severe winters (for example, in the winter of 1939 to 1940) and later, 
as the herd grew, even adult animals, mostly males, died in greater 
numbers under similar winter conditions. Adult males, in the event 
of an acorn shortage, as happened in 1950, could not recoup after the 
period of rut and had to face the hard winter unprepared (Mertts, 
1953 and 1957). 

During the period 1933 to 1952 the winters of 1939 to 1942 and 
1950 to 1952 (in all, five winters) proved particularly hazardous for 
deer of Voronezh presetve. In these winters death due to emaciation 
and other factors (excluding wolves) was established in 113 deer, of 
which 56.6% were juveniles, 41.6% adult males, and 1.8% adult 
females. Adult deer died mainly due to emaciation (instances of 
death became more frequent with the intensification of frosts), while 
juveniles died due to the consumption of large amounts of coarse 
food (Mertts, 1953 and 1957). Later, the deer population in the pre- 
serve rose, but during the extremely snowy and severe winter of 1955 
to 1956 (snow height in some habitats exceeded a meter and the 
minimum temperature fell to-40°C), 177 of the 880 deer died. 
Among deer which died in that winter due to emaciation and exces- 
sive cold, 42.5% were juveniles in their first year and 7.5% in their 
second year, 35% adult males, and 15% adult females. The wastage in 
the group of adult males comprised roughly 27% of the total popula- 
tion, among adult females 6%, and among the current year's brood 
36% (Zharkov, 1957). The death of deer was maximum in sections 
where vegetation had been greatly damaged by cattle in the preced- 
ing summer (Kaznevskii, 1958). In Belovezh Forest starving deer died 
in winter due to consumption of spruce needles, which are unsuita- 
ble to them and cause nephritis and constipation (Wroblewski, 
1927). 

Mass deer mortality due to wolves, emaciation, and in part snow 
avalanches, was observed in the northwest Caucasus in the winters 
of 1910 to 1911 and 1931 to 1932 (Nasimovich, 1938 and 1939). Dur- 
ing the most snowy winters in the last one-third of the nineteenth 
century deer in the Ingur and Tskhenis-Tskala were often totally 
destroyed (Kalinovskii, 1901). Instances of mass mortality of maral 
and Manchurian wapiti in snowy winters were a more rare pheno- 
menon, but even so occurred in some winters, for example that of 
1913 to 1914 (mountains of southern Siberia and the Primor'e 
region; Solov'ev, 1921; Kaplanov, 1948; and others). In the Far East a 
large number of Manchurian wapiti died due to hunters, predators, 
and other reasons in the winter of 1865 to 1866 (Przewalski, 1867) 

"Over 100 more deer died outside the preserve area. 



225 



and in the snowy winter of 1935 to 1936 (Kaplanov, 1948). Mortality 
among maral and Manchurian wapiti due to shortage of adequate 
food was also noted in the winter of 1940 to 1941 (Sayans and Bargu- 
zinsk preserves; Nasimovich, 1955). In Altai preserve mortality of 
maral due to snow avalanches has been recorded (Dul'keit, 1955) and 
in the Primor'e region of Manchurian wapiti due to wolves when 
massive slippery crusts formed on rivers (Okhotina, 1957). 

The winter of 1953 to 1954 was severe in the southern Ukraine 
and accompanied by the formation of ice crusts and deep snow (up 
to 55 cm). In this year, of the 133 deer in Azov-Sivash preserve, 46 
died due to emaciation and excessive cold; of these 42.4% were adult 
males, 36.2% juveniles, and 21.4% adult females (Ishunin, 1956). 
Here, too, male mortality was considerably higher than that of 
166 females. In 1949 unparalleled floods in the Amu-Darya and intense 
flooding in Darganatinsk tugais claimed 50 of about 100 Bukhara 
deer (Klyushkin, 1954). 

While on the subject of deer mortality mention should be made 
of the high mortality rate among adult males in the period of rut. In 
Voronezh preserve over a period of 20 years 27 cases of death of males 
as a result of duels in the period of rut have been recorded, represent- 
ing 13% of all deaths for which reasons have been established 
(Mertts, 1957). The high rate of mortality among adult males iri 
severe winters as a result of emaciation in the period of rut should 
not be overlooked. In Crimean preserve 33 cases of death of deer 
were established for the period 1945 to 1950. Of these, deaths due to 
various traumas, including those inflicted during duels in the 
period of rut, accounted for 30% (Yanushko, 1958). Hence, even in 
those deer populations where their number is uncontrolled, among 
adult animals there is a significant preponderance of females even 
though at birth the ratio of the sexes is close to 1 : 1 (Yanushko, 1958; 
Kroning and Vorreyer, 1957). 

Based on data of encounters between adult male and female deer, 
obviously not very accurate (females usually remain in large groups 
and thus draw immediate attention), the following ratio of males to 
females emerges in different regions: 



Preserve 


Years 


(5:9 




Reference 


Belovezh Forest 


1945-1949 


1 : 1.2 




Sablina, 1955 


Voronezh 


1941-1953 


1 : 1.6 




Mertts, 1953 


Voronezh 


1955 


1 : 1.8 




Zharkov, 1957 


Crimea 


1945-1950 


1 : 1.6 




Yanushko, 1958 


Caucasus 


1927-1935 


1 : 1.7- 


4.0 


Nasimovich, 1936 



226 



Preserve Years б : 9 Reference 



Borzhom up to 1946 1 

Zakatal'sk 1953 1 

Sikhote-Alin 1936-1941 1 



2.5 Z.S. Ekvdmishvili 
3.7 I.F. Popkova 
3.0 Kaplanov, 1948 



A large number of fawns die in the first few weeks or months 
after birth, either because fawning occurred early or the young fell 
victim to predators. Females also die when cold spells recur. Data on 
this subject are extremely scanty, however. In most regions juveniles 
in autumn or early winter populations comprise 11.0 to 13.4% (Nasi- 
movich, 1941; Sablina, 1955; Zharkov, 1957; and others). In Crimean 
preserve, where wolves are absent, the maximum fawn mortality 
takes place at the age of three to four months; of the total number of 
young born in that period about 37% perish, mortality falling 
sharply in subsequent months (Yanushko, 1958). According to the 
calculations of Sablina (1955), based partly on data provided by 
Severtsov (1941), in Belovezh Forest fawn mortality in the first year 
of birth reaches 50% and drops to 40% in the second year (of those 
surviving at the end of the first year). Only 30% of the original 
year-class remains at the end of the second year. 

In various preserves the annual growth of herds varies from 12.5 
to 20.4%, more often 16 to 18% of the total number of animals. The 
highest growth rate of herds has been recorded in certain years in 
Crimean preserve. In the Voronezh, annual growth of the deer popu- 
lation dropped from the end of the 1940's to 12.5% (average for 4 
years) from the former 17.4% (average for 13 years). Merits (1953) 
explains this as due to dispersal of deer beyond the preserve into 
regions where they are greatly threatened by wolves, and also due to 
unfavorable conditions during one winter. The increase in density 
of population and consequent worsening of food sources could also 
perhaps have been contributory factors. 

In taiga regions the most serious competitor for deer is elk; in 
Belovezh Forest, with mixed forests, the ratio of these two species is 
167 more in favor of deer than elk, and the former is far more adaptable 
than all other ungulates with regard to food selection (Vroblevskii, 
1912). The rapid multiplication of deer in Crimean preserve not 
only arrested the population rise of roe deer, but later even caused a 
reduction, perhaps partly due to migration of roe deer into the much 
lower mountain regions falling beyond the limits of the preserve. 
No less than 71 plant species, i.e., over 50% of all known deer food 
plants in that region, are common to both these species (Yanushko, 



227 

1957). In places where true and sika deer coexist the population of 
both is usually insignificant and hence data on competitive relations 
between them are scanty. 

In Crimea beds of red and roe deer are often found in immediate 
proximity of each other; in winter Siberian roe deer are sometimes 
found among herds of Manchurian wapiti (Cherkasov, 1884). On 
encountering other ungulates at a salt lick deer do not descend into 
it until the departure of elk (izyubr' usually avoid salt licks patron- 
ized often by elk) and roe deer await the departure of [true] deer. 

In Voronezh preserve deer visit in winter places where trees have 
been felled by beavers and feed on slender (up to 1.5 cm, aspen) termi- 
nal branches. In Azov-Sivash preserve crows collect in spring the 
wool of molting deer for building their nests, landing for this pur- 
pose directly on their backs (M.D. Shpigov). 

In addition to economic and elemental factors, the population 
dynamics of deer in most parts of the range is also determined by 
man's influence on topography, which alters the original habitat. In 
the Ukraine the human factor over centuries has ultimately led to 
the disappearance of deer there. Bukhara deer have also suffered 
similar consequences as a result of destruction of tugais, and maral 
in Kirgizia have suffered from the cutting of spruce forests and 
reduction in their seedlings (D.P. Dement'ev, 1946). 

Field characteristics. In external appearance true deer are easily 
distinguished from roe deer and sika (see the field characteristics of 
these species). Females of true deer are without antlers and smaller 
in size than males. 

During slow walking the distance between hoofprints is 50 to 70 
cm, increasing during a gallop to 3 to 6 m. Hoofprints are oval and 
slightly elongated. Even on relatively stable soil not only the two 
medial large toes but also the two small lateral ones are usually 
imprinted (impressions of the latter fall in the rear). Deer hoofprints 
are similar to those of wild boar but the two medial toes of deer are 
less mobile and not as widely extensible as in wild boar; the impres- 
sions of the lateral toes of wild boar are more distinct and the hoof- 
prints more elongated. The tracks of males are larger than those of 
females; even a three-year-old male makes a larger or at least equal- 
sized print as an old female. The hoofprint of a female is more 
oblong and narrow than that of a male. The shape of the hoofprint 
of a juvenile is similar to that of a female (young deer walk [rear] 
hoof on [front] hoof or even advance them slightly forward). The 
pace of the male is longer than that of the female; the clearance 
between the imprints of the right and left hooves is also greater in 
the male; the rear footprints of a male are more stunted. The urine 



228 



patch touches the snow between the hoofprints of the fore and rear 
legs in the case of a male and close to the hoofprints of the rear legs 
in the case of a female. 
168 Feces are in the form of oblong, dark cinnamon-colored pellets 
and form a cluster; the pellets are smaller than those of elk but larger 
than those of roe deer. In summer the pellets of an adult male 
measure 3.0 to 4.5 cm x 2.0 cm and of females 3.0 cm x 1.5 cm; the 
pellets of a male are generally more rounded than those of a female 
(the latter's is more elongated). In winter, when deer eat dry food, 
feces are more regular in form; in summer the pellets often fuse and 
their individuality is obliterated; in spring, during transition to 
grass feeding, feces initially take the form of large semifluid pan- 
cakes in which it is difficult to distinguish individual pellets. 




8 



167 Fig. 54. Hoofprint of male deer on soft soil in a forest (reduced). 

Voronezh preserve. October, 1950. Sketch by A.A. Nasimovich. 




168 Fig. 55. Deer feces in spring. Crimean preserve April, 1952. Sketch by 

T.L. Savranskaya. 



229 




Fig. 56. Winter feces of female deer. Voronezh preserve. Sketch by T.L. 
Savranskaya. 



The bed is oval in shape. In winter, while lying down, deer often 
scatter snow with their hooves, but sometimes lie down directly on 
it. Usually they lie with their head in the direction of the entrance. 
When snow is heavy the animals do not lie down at one place for 
long. 

In winter deer groups form characteristic patches measuring up 
to 50 to 70 m in width at places of feeding. From the broken 
branches of deciduous trees and shrubs and the plucked tops of 
young pines these places are recognizable long after the deer have 
fattened. In winter gnawed bark reveals fairly deep furrows, nar- 
rower than those of elk (deer and elk scratch bark with their inci- 
sors); in summer, when bark can be easily peeled from trees, tooth 
marks are not left behind as the bark is removed in whole strips 
(Klyushin, 1949; Formozov, 1952). 

Places of regular habitation of deer reveal well-beaten tracks, 
sometimes running for several kilometers and connecting pasturing 
meadows, salt licks, waterholes, shaded and sunny slopes of moun- 
tains, and so on. The presence of deer can also be judged from 
scratches made on trees by them while attempting to peel the skin 
from their antlers, from shrubs and small trees damaged and dis- 
turbed by the antlers (in such places, the turf and soil are stamped 
down by hooves and there is a distinct smell of urine), from clusters 
of dropped wool adhering to bushes, and so on. 

Frightened deer emit a broken and not very loud "hau" (the cry 
of a frightened male roe deer is far louder). Females and young on 
registering danger emit short and fairly melodious sounds resem- 
bling squeaks; females sometimes make a low mooing sound. In 
nature, however, for most of the year, even in places where deer are 
extremely abundant, their voices are rarely heard. It is only in the 
period of rut that the bellowing of males can be heard over a long 
distance. (A.N.) 



230 

Economic Importance 

Hunting of deer is strictly prohibited in most of their range because 
of their low population, but licensed hunting of adult males is 
permitted in Siberia and Georgia. Deer are killed almost exclusively 
by gun; the time of hunting is usually spring-summer (for velvet 
antlers) and autumn-winter; the best time in the latter period is the 
second half of November and the first half of December. In spring 
169 and early summer deer are caught not so much for their meat (spring 
deer are often lean), as for obtaining young unossified antlers — the 
velvet antlers used for medicinal purposes (to enhance overall vital- 
ity and so on). Velvet antlers are of maximum value when they are 
not fully developed (this corresponds in different parts of the range 
roughly to May 15 to 25 through June 15 to 25). For obtaining velvet 
antlers deer are caught at salt licks, banks of water sources, well- 
frequented meadows, during transit, etc. Sometimes dogs are used to 
bring deer to bay; dogs are also used to track wounded animals. In 
winter deer are caught on the snow most often by stealth but some- 
times by organizing a beat. Occasionally dogs are used and in this 
case deer run into rocky areas and stand there in sheltered spots; 
hunters take advantage of this habit. 

Until recently hunting was widely practiced during the period of 
the rut; by locating the direction of bellowing the hunter 
approached the deer within gun range. To entice the stag to bellow 
more often, its bellow was imitated by cupping the hands in front of 
the mouth or a pipe made from stalks of cow parsnip, birch bark, 
etc. was used (hunting "with a pipe"). Many males during the 
period of rut are greatly emaciated; their meat takes on a bluish tinge 
and has an unpleasant taste. The products of such a hunt are of low 
value except for the hide, which at that time is of good quality. From 
the sportsman's point of view hunting "by bellowing" is one of the 
most interesting sports. 

In pre-Revolutionary Russia a very large number of deer were 
caught by chasing them on ice crust or in deep snow. Deer were also 
caught in deep snow to replenish maral nurseries but a large 
number became exhausted in this process and died. At some places 
in Siberia such a catch resulted in greater loss of life and affected the 
population of deer more than ordinary hunting. Now the chasing of 
deer on ice crust and in deep snow is prohibited. 

Also prohibited at present is the trapping of deer, extensively 
practiced before, mostly in Siberia and the Far East. Special, long 

'"^Nevertheless poachers even today sometimes use snares made of cable to catch 
deer. 



231 

fences were built with pits dug at exits. Traps were also laid on deer 
trails and at salt licks. 

In some southern regions of Siberia deer were formerly caught in 
the course of their autumn-winter migrations and near points of 
river crossings. This type of hunting is practiced very rarely 
nowadays. 

For catching deer alive different types of traps are used (corrals, 
mobile traps with closing shutters, bait, etc.), but none is very effec- 
tive. In Voronezh preserve chasing on snow was done to capture deer 
alive and nets sometimes employed. 

By the end of the nineteenth century up to 25,000 or more deer 
had been caught in Siberia and the Far East (Solov'ev, 1922; and 
others). The main products of deer hunting are meat, hide, and 
velvet and ossified antlers; secondary products include tails, sex 
organs of males ("pyrki"), and fetuses removed from wombs 
("lutai"). Secondary products are used in some regions of the USSR 
(Evenk, Nanaians, and others) as a stimulant during child-birth and 
for the preparation of drugs for different ailments (heart and sex 
disorders, malaria, and others). The quality of deer meat is excel- 
lent in autumn before the onset of rut. At places where food is 
abundantly available and males are able to recoup at the end of rut, 
their meat as well as that of females is of high quality during the 
entire first half of winter. The meat yield, i.e., the body weight 
without head, hide, viscera, and hooves, constitutes 55 to 65% of the 
animal weight. A large maral (see section on growth and develop- 
ment) can yield up to 220 to 230 kg of meat (including up to 15 to 25 
kg fat), a Manchurian wapiti up to 190 to 200 kg, a Voronezh deer 
rarely over 150 to 160 kg, and a Crimean deer generally less than 140 
kg. 
170 The high value of maral and Manchurian wapiti velvet antlers 
(less, however, than that of sika) led even in the nineteenth century 
to the founding of a special branch of animal husbandry, i.e., catch- 
ing and raising of deer in paddocks. In the Altai raising maral 
commenced in the 30s of the nineteenth century on the Bukhtarma 
River. By the end of the century there were 200 nurseries established 
in the Altai, containing over 3,000 maral. In Trans-Baikal (along the 
Ingoda River) raising Manchurian wapiti commenced in 1843 and 
by the end of the nineteenth century there were 300 farms, contain- 
ing over 1,000 wapiti (Turkin and Satunin, 1902). The ^xtent of 
maral farming in the past in the Altai and the hunting of deer can 
also be judged from the fact that present-day Gorno-Altai Autonom- 

" For more details, see Skalon et al., 1951 and Abramov, 1954. 



232 

ous Region prepared 6,600 maral hides and 1,116 pairs of velvet 
antlers in 1896 (Yukhnev, 1903). 

Raising maral is now most extensively developed in the Altai. A 
valuable endocrine preparation, pantocrine, is prepared from the 
velvet antlers of maral and Manchurian wapiti (formerly the velvet 
antlers were mainly exported to China). European red deer have 
never been a source of velvet antlers. 

In pre-Revolutionary years ossified antlers were exported to 
China (used in Tibetan medicine) and also to western Europe as 
decorative trophies. For the latter purpose antlers were removed 
along with part of the frontal bone. Even in the early twentieth 
century antler buyers from various cities, even Paris, assembled in 
the Krasnodar region. Antlers were used in making knife handles 
and other articles and for making glue by boiling them. 

The quality of deer hide is best in the latter half of September to 
October. Hides are converted into tough chamois, and the hide with 
hair converted into warm fur coats, aviator boots, mittens, and other 
articles of apparel, covers for yurts (nomad tents), and edges for skis. 
Autumn hides, which are the toughest, are used for belts, breeches, 
bridles, etc.; spring hides, because of holes made by gadflies, are less 
valuable. 

In the distant past attempts were made in Poland and Germany 
to use deer as draft animals (Turkin and Satunin, 1902). 

Acclimatization of deer has long been attempted. In Belovezh 
Forest, where deer were extinct even by the middle of the nineteenth 
century, 18 deer were imported from Silesia in 1864. Later, from 1891 
to 1902, about 300 deer were imported from Silesia, Bohemia, and 
Spala. During the 1870's and 1880's deer were imported from Poland 
or Germany into Ramon' and held captive until 1917 when they 
were set free. These deer represent the parent stock of the present-day 
Voronezh animals. In setting up the Borzhom game farm 83 deer 
had been imported by the end of the nineteenth century, mostly from 
Austria (58) and later from some regions of the Caucasus and 
Vil'nyus (in the latter case deer were not of local origin). The intro- 
duced deer interbred with native animals (Anon., 1891) and hence 
the current population of Borzhom deer is of hybrid origin. In pre- 
Revolutionary years deer were also released in the Baltic (imported 
from Belovezh Forest and western Europe), western Ukraine 
(imported from western Europe) and Askaniya-Nova (for the first 
time around 1894). In the latter region Crimean deer, Altai maral, 
and complex hybrids sharing the blood of several deer forms, includ- 
ing sika, can be seen today. In Latvia deer were reacclimatized not 
long before World War I (Kalnin'sh, 1950). 



233 

A large number of experiments for acclimatization of deer have 
been conducted in the last few decades. In 1928 five deer of hybrid 
origin were imported from Askaniya-Nova into the Biryuchii 
Islands (Black Sea). Nine more deer were added in 1946, also from 
Askaniya-Nova, and three in 1951 (V.A. Mattskov). In 1929 twenty- 
six hybrid deer from Askaniya-Nova were released in the region of 
Pecheneg village in Kharkov district. The deer released there have 
survived (in 1948 there were 16 animals, which rose to 60 before the 
Great Fatherland War [W.W. II]; N.V. Charlemagne). In the early 
1950's ten deer were imported from Crimean preserve and twelve 
from Voronezh into Pechenezhsk game farm. Deer were also released 
in Gavrilov game farm of Kherson district, Ukraine, where by the 
end of the 1940's over fifty animals were counted. As a result of the 
construction of a water reservoir there almost no deer are seen now. 
In 1957 sixteen Voronezh deer were released in Korostansk forest 
farm of Zhitomir district, twenty-four (also from Voronezh preserve) 
in Vovchansk forest farm of Kharkov district, and sixteen from Cri- 
mean preserve inuoduced in Brovar region of Kiev district (A. P. 
Korneev). 

After the First World War deer were introduced from Germany 
into Estonia and raised under conditions of partial confinement on 
the Abruk Islands (near Saaremaa Island) and northwest of Pyarnsk 
(Audi Forest). Only a few have survived in the latter region to date. 
In 1957 eight maral were introduced into Estonia and placed for the 
first time in confinement (E.V. Kumari). Also in 1957 thirty-seven 
deer were brought from Voronezh preserve into the Lithuanian 
Soviet Socialist Republic. In 1929 six European deer were brought 
into the territory of the present Molodechnensk district of the Belo- 
russian Soviet Socialist Republic, then part of Poland. These deer 
formed the nucleus of a small population, surviving even at present 
in Ivenets and Volozhinsk regions. In 1949 they numbered fifty 
(Serzhanin, 1955). In 1957 forty-eight deer from Voronezh preserve 
were brought into Berezinsk beaver preserve (Nemansk basin) and 
thirty-six introduced into Krasnyi Les in 1948 (Krasnodar region). 
From 1940 to 1941 maral from Shebalinsk maral farm (Altai) were 
brought into Bashkir preserve and fifty-three set free in 1941 to 1942. 
They multiplied and later settled outside the limits of the preserve 
(Kaznevskii, 1954 and 1956; and others). In the same period Askaniya 
maral (complex hybrid) were acclimatized in Mordov preserve. 

A few Caucasian deer were held for several years in Teberdin 
preserve. In 1953 four deer from that nursery were released in the 
Gonachkhir River region (deer in Teberda region became extinct 



234 

even in the nineteenth century). In 1955 there were eleven deer (Inya- 
kova, 1957). 

In Zavidov game farm (north of Moscow), eight maral were 
released in 1938 but during the Great Fatherland War all were des- 
troyed (Danilov, 1947). Recently, no less than seventy maral from 
Altai maral farms have been released in batches of six to thirteen 
animals in several game farms in Moscow, Kalininsk, and Yaroslav 
regions (Yurgenson, 1957; and others). 

The main drawback in most cases of release of deer is the un- 
acceptability of hybrid forms and release in the same region of deer 
belonging to different subspecies. In this context the preservation of 
Crimean deer, which are an entirely distinct subspecies, in pure 
form without admixture with other forms, should be safeguarded.'^^ 

One of the important problems of present-day game and sanctu- 
ary farming is to combine raising and protection of deer with affo- 
restation. The experience of Belovezh Forest, Voronezh, and 
Crimean preserves, the Baltic republics, and western European game 
farms has shown that when deer population density is high, the 
animals adversely affect forest economy. Even though they only 
damage seedlings and shrubs severely in some winters, this renders 
the raising of mixed plantations by selecting deciduous seedlings 
and weeding out coniferous species difficult. A progressive increase 
in spruce and a decrease in proportion of deciduous species in the 
forest composition of Belovezh Forest was observed in the early 
twentieth century and associated with the then-high deer popula- 
tion (Vrublevskii, 1912; Kalnin'sh, 1950; Bannikov and Lebedova, 
1956; Yanushko, 1957; and others). 

In Voronezh preserve deer at places damage saplings of oak, 
pine, and other plants; in Belovezh Forest they damage saplings of 
ash, maple, oak, and other plants, while in Crimean preserve, dam- 
age is mostly confined to saplings of oak, beech, pine, maple, and 
some shrubs. In the Primor'e region Manchurian wapiti greatly 
damage philodendron and aralia. In places where efforts are made to 
combine forest growth and raising of deer the density of the latter 
should not exceed an average of 10 per 1,000 hectares of land; in 
sections of the range where food availability is less, this norm should 
evidently be reduced and in the south, where winter is not very 
severe, it should be raised slightly. At the time of implementing 
various forestry measures, shrubs and saplings should be preserved 
since they constitute the main food of deer in the winter. It should 

A project for acclimatization of deer from other parts of the range in the 
mountain forests of Crimea was proposed in 1958. 



235 

also be remembered that when the population density of deer goes 
up, the animals and their antlers are reduced in size, their fertility 
decreases, and many other undesirable consequences follow (Caesar, 
1956; and others). 

In some instances forest deer, when relatively abundant, can 
threaten adjoining agricultural farms, kitchen gardens, melon 
fields, and haystacks left in forests. In the Crimea deer at places 
damage vineyards (they eat the fruit, break branches, and trample 
bushes) and visit fruit gardens. In the Amu-Darya tugais Bukhara 
deer visit farms where they avidly consume the herbage of mungo 
beans {Phaseolus mungo), damaging the plant along with flowers 
and green pods. Joughara [Sorghum cernum,^, sesame, lucerne 
[alfalfa], and muskmelon are also sometimes damaged by deer 
(Klyushkin, 1949). On the whole, however, these losses are rather 
insignificant and can easily be prevented by implementmg appro- 
priate measures for the control of deer populations. (A.N.) 

Subfamily of Elk, or Telometacarpal Deer 

Subfamily Odocoileinae Pocock, 1923 

Deer of small, moderate, or large size. 

Only distal extremities of metacarpals retained while proximal 
portions reduced to insignificant remnants, usually fused with can- 
non bone. Cuneiform either separate from naviculo-cuboid or fused 
with it. 

Depression for preorbital gland on facial part of lacrimal, with a 
few exceptions {Capreolus and one species of Pudu), present and 
well developed. In several cases vomer divides choanae. Antlers inva- 
riably present; in one case (Rangifer) even females have antlers. 
Antlers extremely diverse in form and develop from simple, 
unbranched, short spike to very complex designs. Brow tine usually 
absent and later half of main beam often curves forward. 

Tail short, of average length, or long. Bare patch at end of muz- 
zle variable; sometimes very much reduced, absent, or very promi- 
nent. With a few exceptions (Capreolus and one species of Pudu), 
preorbital gland present and well developed. Toe pads highly devel- 
oped and cover entire lower surface of hoof. Preputial portion of 
penis, with a few exceptions, fused with abdominal skin. In other 
features Odocoileinae do not differ from Cervinae. 

These are forest, forest-steppe, and tundra animals. 

Odocoileinae represent a typical group of deer of the New 
World, where they are widely scattered, very diverse,^and represented 



237 

by a maximum number of genera and species. In the Old World only 
one genus (Capreolus) and one species of the subfamily are found. 
Two genera {Alces and Rangifer) are present in both the New and 
Old Worlds. In the New World Odocoileinae cover the entire range 
of the family; in Eurasia, they are distributed in the north up to the 
northern boundary of the family and in the south to the Mediterra- 
nean Sea, Asia Minor, northern Iraq, Ferghana, northern Mongolia, 
northeast China, Kansu, Szechwan, and Cheju-Do Island (for more 
details, see description of distribution of roe deer Capreolus 
capreolus). 

The subfamily is well distinguished from all other subfamilies of 
Cervidae, but very close to subfamily Cervinae and even sometimes 
included in it. 

Odocoileinae are known from the Lower Pliocene of Europe and 
Asia (Capreolini). In the New World they have been detected from 
the Pliocene. 

Odocoileinae is understood here in its more or less traditional 

174 format (according to Simpson, 1945, with the exclusion of genus 

Hydropotes from the subfamily), and not in the sense of Neocervi- 

nae Carette, 1922 (all Odocoileinae except Alces and Capreolus; 

Flerov, 1952). 

The systematics of the group, mainly of South American forms, 
has attracted attention recently and their species composition and 
generic groupings, which until recently remained totally vague, 
have been delineated. Nevertheless differing opinions still persist. 
Simpson (1945) grouped the species (not inchxding Hydropotes) into 
nine genera, Haltenorth and Trenze (1957) into seven, and Flerov 
(1952) into six. These differences are mainly due to downgrading 
some genera to the rank of a subgenus or simply discarding them. As 
far as the number of species is concerned, there are 15 according to 
the latest concepts (Haltenorth and Trenze, 1957), adopted in the 
present work (23 according to Flerov, 1952), which constitute about 
42% of the species of the subfamily. In all, the subfamily comprises 
12 genera (24% of all genera of the family) of which 7 are recent (41% 
of surviving genera of the family) and 5 extinct. Extant genera are: 
Odocoileus (subgenera Odocoileus, Blastocerus, and Dorcelaphus: 
four species); Hippocamelus (two species); Mazama (four species ); 
Pudu (subgenera Pudu and Pudella: two species); Capreolus (one 
species); Alces (one species); and Rangifer (one species). 

The first four genera form a single, fairly intimate group (tribe) 
of purely American forms, and three other separate, sharply isolated 
monotypic suprageneric groups (tribes). Capreolus is the lone form 
of the subfamily found in Eurasia, while Alces and Rangifer are 



238 

encountered in the Old and New Worlds. 

In the USSR three genera occur (about 43% of genera of the 
subfamily): reindeer — Rangifer, elk — A Ices, and roe deer — Capreolus, 
with three species (20% of species of the subfamily). They constitute 
about 1.0% of the species of Russian fauna. 

In the USSR Odocoileinae are distributed in tundra and forest 
zones, Crimean and Caucasian forests, and the mountains in the 
eastern part of Central Asia. The geographic range and population 
of one species decreased sharply in the last century, while the range 
of the others remained quite unchanged or only insignificantly 
altered. 

Odocoileinae represent important game and table animals which 
are often protected. One species (reindeer) survives in both wild and 
domesticated states. (V.H.) 

Genus of Roe Deer 

Genus Capreolus Gray, 1821 

1775. Capreolus. Frisch. Natursyst. d. vierfdss. Thiere, p. 3. This 

name, as well as all generic names given in this compilation 

(except Dama) are unsatisfactory from the viewpoint of 

nomenclature. 

1821. Capreolus. Gray. London Med. Report, vol. 15, p. 307. Cervus 

capreolus Linnaeus, 1758. 
1837. Caprea. Ogilby P.Z.S.L., 1836, p. 135. Cervus capreolus 
Linnaeus, 1758. (V.H.) 
Capreolus are telometacarpal deer of small size, very light and 
slender build, with a relatively short trunk, long neck, straight 
back, relatively small head, and long legs. 

Hooves of middle toes small, narrow, and sharp; lateral hooves 
small, set high, and nonfunctional. 

Skull relatively small, with a somewhat shortened facial portion. 
Nasals normal in length (roughly equal to length of upper tooth 
row; nasal and premaxilla not more than length of upper tooth 
row). Depression absent on anterior part of premaxilla. Nasal pro- 
cesses of premaxillae reach, or almost reach, nasals. Preorbital 
depression on lacrimal weakly developed, small, and with blurred 
outline. Ethmoid pit well developed. Processes for horns (or cornu) 
on frontals set back and up, and close together. Rear portion of 
vomer low and does not interrupt choanae. 

Upper canines usually absent both in males and females, but 
seen exceptionally in males. Upper molars with relatively low 



239 

crowns without supplemental columns on internal side. Dental 
formula: 

.00 3 3 „g 

1-, C-, pm-, m^, = 32. 

Only males sport antlers, which are very small. Maximum length 
of antlers not more than 1.5 to 2.0 times length of head; however, 
often shorter than head. Antlers sometimes develop in very old 
females but are invariably small and usually irregular in form. 
Antler beam round in section and almost straight or only slightly 
curved backward at end; usually with three tines at terminal portion: 
one set forward, one upward, and one backward. Exceptionally 
rear tine bifurcate (four tines on antlers) and, even more rarely, 
upper tine also (five tines on antlers). Antler beam with uneven 
surface, particularly rough on inner surface of the bottom half or 
third of antler length. Sharp tuberosities and often small elongated 
prominences form there. Antlers turn upward and backward and 
appear to be set almost vertically. Beam bases usually close-set and 
beams run almost parallel; sometimes slight or significant spread of 
antlers seen. Often they form a distinct layrate figure. Rosette rela- 
tively well developed. 

Ears long and attenuate. Tail rudimentary and does not project 
out of fur and imperceptible in a living roe deer. Muzzle somewhat 
stunted and blunted and tip bare; bare field large and encompasses 
nostrils. Dewlap absent. Preorbital glands rudimentary and do not 
function. Hoof glands present only in hind limbs, in the form of a 
saccate invagination with a small opening. Tarsal glands well 
developed. 

Coloration uniform; in summer coat bright and rusty and in 
winter dull and gray. Legs of same color as trunk. Large bright 
patch (speculum) occurs on rear side of thigh and around tail. Mane 
on neck absent. Coloration of young deer (in first coat) spotted. 
Female somewhat smaller than male. Teats four. 

These animals inhabit deciduous and mixed forests in the plains 
and mountains, southern borders of tugais, forest-steppes, river val- 
leys in the steppes, and sometimes even reeds. They are encountered 
in mountains up to heights of 3,000 m above sea level. 

The distribution of the genus is restricted to Eurasia. Its geogra- 
phic range (reconstructed) used to cover western, southern, and cen- 



'^'Front tine usually considered medial, while upper one represents tip of the beam; 
rear tine set backward. 



240 




176 Fig. 58. Reconstructed geographic range of roe deer genus (Capreolus Gray) and 
species Capreolus capreolus L. V.G. Heptner. 



tral Europe, including England/^° Italy, and Sicily (absent in 
remaining Mediterranean Islands), and in the north up to the south- 
ern and central parts of the Scandinavian Peninsula, Asia Minor, 
Syrian region of the United Arab Republic and Palestine, northern 
Iraq, northern (Elbruz) Iran, the Caucasus, and Crimea. 

In the east European plains and Asia the northern boundary ran 
along the Gulf of Finland, slightly south of Moscow, through the 
mouth of the Kama and the upper reaches of the Pechora, through 
Tobol'sk, Novosibirsk, the mouth of the Angara (upper Tunguska), 
and the northern extremity of Baikal. The eastern boundary rose up 
to 60° N lat. and, descending south, intersected the lower courses of 
the Amur and ran toward Tatarskiy Strait. The southern range 
covered the Korean Peninsula, northeast China, northern parts of 
176 Mongolia (Pri-Hobsogol and Khentei) and the northern and north- 
western parts of eastern and southeastern China (Jehol, Chakhar [?], 
Hopei, Shansi, Shensi, Kansu, and northern Szechwan). The west- 
ern range encompassed the Altai range, Semireche, the Tien Shan 



Contrary to reports by some authors (Flerov, 1952), apparently roe deer never 
occurred in Ireland. 



241 

mountain system up to Ferghana valley, and Kazakhstan (northern 
half). 

In general the typical zone of the temperate Eurasian belt repre- 
sented the range of the genus. Compared with the reconstructed 
range described earlier, the actual present-day range at places has 
shrunk quite significantly or even broken down into individual 
pockets (see "Geographic Distribution" under roe deer). 

The genus Capreolus stands very sharply isolated within the 
subfamily and exhibits no apparent close association with other 
present-day genera. 

This situation is largely explained by Capreolus being relatively 
ancient, one of the very oldest living genera of the family. Forms 
placed in this genus or proximate to it are already known in the 
Middle Pliocene (Europe). Genus Procapreolus, which is very 
close to Capreolus, is known from the Lower Pliocene (according to 
some authors, even from the Upper Miocene) of Europe and Asia. 
There is reason to consider this group the parent stock of 
Capreolus. 

The distribution of fossil forms of the genus (reconstructed 
range) generally does not fall significantly beyond the limits of the 
present-day range. Species still different from extant species lived in 
the Gunz period and extant species appeared in the Riss-Wurm 
period (Flerov, 1950). 

Three species {capreolus, pygargus, and bed for di) were recog- 
nized in the genus until recently. In reality, however, only one spe- 
cies exists with some geographic variations (about 5% of species of 
178 the family). The differences between some, especially between 
capreolus and pygargus, are so marked that they earlier prompted 
recognition of two "species" in the genus. 

These are game and table animals caught mainly for their meat 
and hide. The antlers are used as decorative trophies and for making 
various articles. The animals are shot for sport and raised in con- 
trolled game farms. 



The view that Capreolus undoubtedly represent a connecting link between the 
groups Muntjacus and Elaphodus (Cervulinae according to some) and red deer 
(Cervinae) (Flerov, 1952) is hardly justifiable. 

'^^Some researchers date fossil finds of true Capreolus to a much later period. 

Those who include Capreolus (along with Alces) in the genus of red deer 
(Cervinae) and separate this genus from American deer ("Neocervinae") place in a 
series the forms leading to Capreolus and Pliocervus (particularly Flerov, 1952) 
belonging to Cervinae. In any case the antiquity of the independent existence of this 
genus is beyond dispute. 



242 




177 Fig. 59. Siberian roe deer, Capreolus capreolus pygargiis Pall. Sketch by A.N. 

Komarov. 



243 

One species, Capreolus capreolus Linnaeus, 1758, is found in the 
USSR and represents about 0.3% of all species of Russian fauna. 

Roe deer represent table animals which at places are of great 
interest as objects of sport. Hunting them is prohibited in some 
parts of the range. (V.H.) 

5. ROE DEER '^' 

Capreolus capreolus Linnaeus, 1758 

1758. Cervus capreolus. Linnaeus. Syst. Nat., 10th ed., vol. 1, p. 68 

Sweden. 
mS. Cervus pygargus. Pallas. Reise, vol. 1, p. 453. Sok River, left 

tributary of Volga, former Samar province. Locality fixed as 

source of Sok, i.e., Bugulma-Belebey rise.^^^ 
1832. Capreolus vulgaris. Fitzinger. Beitr. z. Landesk. Osterreichs, 

vol. 1, p. 317. Austria. 
\84Ъ. Capreolus caprea. Gray. List Mamm. Brit. Mus., p. 176. 

Proposed as substitute for Capreolus Linn. 
184:6. Capreolus europaeus. Sundevall. K. Svenska Vetensk. Akad. 

Handl., 1844, p. 184. Proposed as a substitute for capreolus 

Linn. 
1889. Cervus pygargus mantschuricus. Noack. Humboldt., vol. 8, p. 

9. Preoccupied by С mantschuricus Swinhoe, 1864. 
\9^b. Capreolus tianschanicus. Satunin. Zool. Anz., vol. 30, p. 527. 

Kuldja, Tien Shan. 
1907. Capreolus transsylvanicus. Matschie Weidwerk in Wort und 

Bild., vol. 16, p. 224. Banat, Rumania. 

""Roe deer have also been labeled "wild goat" or simply "goat". In Russia the male 
is called "kozel" and in Siberia "iguran"; the female is called "koza" and "kozlukha". 
"Kosulya" is evidently a variation of "kozulya". It has also been suggested that the 
name is derived from the slanted pupils of the eyes (Flerov, 1952); this interpretation of 
the origin of the term begs credulity. 

'"Sometimes the synonyms of this species include (Flerov, 1952) Cervus ahu as 
referenced by S.G. Gmelin (Reise durch Russland, vol. 3, p. 496, 1780). This name, 
applied to northern Iranian roe deer, could have had some nomenclatural significance 
as one of the earliest names except for a simple misunderstanding. The point here is 
that Graelin's work does not mention the name Cervus ahu at all, either on the page 
referenced above or at any other place. On p. 496, in fact, the word "ahu" occurs in the 
margin but is cited as a local name and set in Gothic type. On pp. 496-499 Gmelin 
describes some animal which he had with him, distinctly a roe deer, and presents a 
drawing(Plate 56) but does not give it a scientific title. At the end of the text (p. 499) he 
states that Pallas, to whom a description of the animal was sent, wrote that it was an 
"ahu," evidently Cervus pygargus. Lydekker (1915), who used the name Сегош ahu, 
was evidently responsible for its introduction into nomenclature. 



244 

1908. Capreolus bedfordi. Thomas. Abstr. P.Z.S.L., vol. 32, p. 645. 
Chaochen-Shan mountains 100 miles west-northwest of Tai- 
yuan fu, Shansi province. 

1909. Capreolus pygargus var. ferganicus. Rasevig, Sem'ya Okhotnikov, 
no. 8, p. 160. Precise location not indicated but evidently 
Ferghana. Mountains adjoining Ferghana valley from the 
north have been suggested. 

1910. Capreolus capreolus balticus. Matschie. Weidwerk in Wort 
und Bild., vol. 19, p. 263. "Vikherstgof, eastern Prussia." 

1919. Capreolus capreolus albicus. Matschie. Weidwerk in Wort und 

Bild., vol. 19. p. 263. Ezerki near Lissa, Poland. 
1910. Capreolus pygargus Pall. var. caucica. Dinnik. Animals of the 

Caucasus, vol. 1, p. 73. Northern Caucasus. 
179 1912. Capreolus capreolus warthae.MsLischie. Deutsche Jagerzeitung, 

vol. 58, p. 801. Dombrovo to east of Bytom (Beiten), Poland. 
1916. Capreolus capreolus armenius. Blackler. Ann. Mag. Nat. Hist., 

vol. 18, p. 78. Sumela, near Trabzon, Asia Minor. 
1916. Capreolus zedlitzi, Matschie, Sitzungsber. Ges. Naturf. Freunde 

Berlin, p. 172. Slonim to southwest of Grodno, Belorussia. 
1935. Capreolus capreolus ochracea. Barclay. Ann. Mag. Nat. Hist., 

vol. 15, p. 627. Korean Peninsula. (V.H.) 

Diagnosis 

The genus contains only one species. 

Description 

Size small, second only to musk deer from among Russian species of 
the family, and smallest of Russian deer. 

Build extremely light and slender, trunk relatively short, and legs 
long. Neck quite long; that of males thicker than that of females. 
Head relatively short, highly wedge-shaped, and pointed in front. 
Back straight, hind legs somewhat longer than forelegs, and croup 
markedly higher than withers (design of jumping animals). Ears 
fairly large, long (longer than one-half of head length), and rela- 
tively broad. Bare patch at end of muzzle large, rising above the 
upper edge of nostrils, and surrounding them from the outside. Tail 
very short, does not project out of fur, and not visible externally. 
Preorbital gland poorly developed; hoof glands present; metatarsal 
glands well developed, with tufts of long, light-colored hair. Cen- 
tral hooves small and narrow and lateral hooves tiny and set high. 

There are two main types of antlers, differing in structural char- 



245 

acteristics but mainly in size.'^^ Antlers of the European type are 
small, do not, or only slightly, exceed skull length (length up to 25, 
rarely 30 cm), and never exceed head length. An additional tine is 
very rare; usually only three tines present. Beams set very straight, 
sometimes almost parallel, or with a slight lyrate flexure. All tines 
usually set in the same plane. Bases of antlers close-set; rosettes large 
in diameter, adjoin or almost adjoin each other, and often closely 
pressed to each other. Distance between bases of beams usually not 
more than diameter of beam at that height. Osseous growths [pedi- 
cels] ("pipes") on frontals close-set, usually parallel, and sometimes 
tips quite close. 

Tuberosities on beams developed mainly up to first tine; they are 
larger and coarser along inner and posterior surfaces of antlers. On 
posterior surface they grow up to the base of the last (third) tine. 
Tuberosities are well manifested but relatively small and usually do 
not exceed 1 cm in height even on large antlers. 

Antlers of the Siberian type are large and, when fully developed, 
not less than length of skull, or longer, or even markedly longer. 
Antler length along main beam may reach 45 cm (or even more). 
Usually antlers of this type represent a simple but magnified replica 
of the European type with the difference that the osseous growths on 
the frontals are relatively more wide-set, their ends usually diverge, 
the rosettes are relatively small and do not touch each other, the 
antler bases are wide-set, and the distance between them is more than 
the beam diameter. Quite often even the distance between the 
rosettes is more than the antler diameter. Tuberosities are also more 
prominent. 

In an extreme variation the Siberian type of antlers may be 
wide-set, the rosettes not large, and the distance between them not 
less than the antler diameter. The beams diverge sideways right from 
180 the base. Often the beams are fairly straight and sometimes even 
form a lyrate figure. The antler tip often curves sharply inside; the 
rear tine also turns inside and quite often bifurcates. Even the for- 
ward tines reveal a tendency to bend in the same direction. The 
antler beam and tines are massive. The tuberosities on the beam are 
fewer than on the European type but more massive and prominent, 
their height often exceeding 2 cm. Some look like small irregular 
additional tines. 

Sometimes not only the rear tines, but also the second ones (top 
of beam), are bifurcate and thus five tines occur. Sometimes the 

Innumerable abnormalities in development of antlers described extensively in 
game literature have not been included here. They are relatively rarely seen in the 
Soviet Union and occur mainly in central and western Europe where living condi- 
tions of roe deer differ significantly from those of the LT5SR. 



246 

number of tines is even greater (up to 22), but this is purely a patho- 
logical phenomenon. Another type of pathological alteration also 
occurs, namely, an antler in the shape of a "wig", i.e., a solid osse- 
ous mass in the form of an "accumulation" covering the head and 
even the eyes. Small irregular antlers are occasionally seen in old or 
abnormal females. In general, among antlers of the Siberian type the 
same abnormal deviations as found among European forms are 
seen, but relatively rarely. 

The normal forms of the two antler types differ very sharply, 
providing a basis for the earlier division of roe deer into two species. 
At the same time the Siberian type of antlers differ in different parts 
of the range and there are also populations in which antler develop- 
ment is of an intermediate nature. 

Full antler development is seen in roe deer older than three years. 
In fawns of the first year only tiny prominences ("pipes") are seen by 
autumn, simple unbranched "rods" in the second year, two tines 
after the second year, and three tines after the third. Later, normally, 
only an increase in size and weight of antlers occurs. Maximum 
antler development has been recorded at ages five to eight years. 
Deterioration sets in later and an overall reduction in size, simplifi- 
cation of design etc., associated with old age, is normal (roe deer live 
up to 15 years in captivity, but rarely beyond 10 years in nature). 

The coloration of adult animals is fairly uniform. Only a large 
white or yellowish-white speculum stands out prominently in the 
caudal region, the width of which can reach 25 to 30 cm. At the top 
181 the speculum does not spread above the caudal base. Its width is 
usually more than its height; sometimes it is nearly an inverted, 
heart-shaped form. The general shade of the winter coat of the trunk 
is gray, which shows some geographic as well as individual variabil- 
ity; it may be light gray or bluish-gray, or, contrarily, darker after 
acquiring a brown or cinnamon tinge. On the rear, along the upper 
border of the speculum, usually a dark band sharply demarcates the 
speculum from the back. The groins, abdomen, and inner sides of 
the limbs are whitish, often with a yellowish tinge. The outer sur- 
face of the limbs is generally of the same color as the trunk but turns 
slightly rusty downward. 

The neck and head are generally of the same color as the trunk. 
There is often (especially in Siberian males) a light-colored patch of 
irregular shape on the throat. The ears are of the same color as the 
head on the outside, but often darker toward the tips; they are whit- 
ish or rusty on the inner side. A white spot of variable dimensions 

Additional data on antler variations in Siberian roe deer (diagrams) are available 
in Fetisov's work (1953). 




247 



Fig. 60. Antler types of roe deer inhabiting the USSR. 

Top row — European roe deer from Carpathians (C. c. capreolus Linn.); others — 

Siberian roe deer from Pri-Baikal (C. с pygargus Pall.). Original drawing, based on 

material from V.G. Heptner. 

occurs on the chin and a dark spot on the lower lip toward the 
corner of the mouth. The upper lip is somewhat darker in color than 
the rest of the head. The winter hair coat consists of underfur and 
long (up to 55 to 65 cm) top and guard hair. The top hair is very 
С1дг1у, thick, and extremely brittle, due to which the coat is easily 
worn out and damaged. This is associated, as in some other deer, 



248 

with the presence in the hair of large air-filled cavities (heat insula- 
tion). The hair is longer on the trunk, especially in the rear, shorter 
on the neck and head, and particularly short on the limbs. A mane 
and elongated hairs below the neck are invariably absent. 

The summer coat is bright, rust-colored or rusty-red, being 
darker, fuller, and sometimes brownish on the spine. The trunk 
coloration lightens downward. The outer surface of the limbs is 
rust-colored, while the abdomen, groins, arm pit region, and the 
inner surface of the limbs are ocherous-white. The head and neck are 
generally of the same color as the trunk, the color of the head corre- 
sponding to that of the winter coat. Sometimes a white or light- 
colored spot occurs on the upper lip. The speculum is less 
prominent and has a yellowish tinge. The hair is much shorter than 
in the winter and is not longer than 35 mm in the rear of the trunk. 
On the head it is much shorter (on the forehead, it is somewhat 
longer, softer, and curly); the hair on the limbs is particularly short, 
only 12 or 13 mm long. 

There are no sex-related differences in the winter or summer 
coat. 

The main shade of color on the upper surface of young animals 
in the first few weeks after birth is rusty-yellow or rusty-cinnamon. 
The abdomen and the inner portions of the limbs are light ocherous. 
The color on the top of the head generally corresponds to that on the 
top of the body, the lower surface being somewhat lighter. Numer- 
ous white or slightly ocherous spots are scattered along the back and 
sides. They are usually so arranged as to form distinct patterns or 
bands. The two central bands are brighter, commence on the occiput 
behind the ears, and extend along the spine right up to the tail. 
Below them, along the sides, usually one or two more such bands 
commence from the shoulder blades and extend right up to the rear 
of the thighs. They are usually less distinct and the spots in them 
less sharp. On the thighs and partly on the shoulders spots are seen 
lower down. Young animals sport this coat for a short while and by 
age four months usually change over to an adult coat of uniform 
color. 

Skull short and relatively broad; zygomatic width roughly about 
one-half length of skull. Facial part of skull short; line dividing 
skull into two fairly equal anterior and posterior portions runs 
roughly through midorbit. Premaxillae short, shorter than upper 
row of teeth, and taper gradually toward end; nasal processes come 
close to nasals or even up to them, but are not wedged between 
nasals and premaxillae. Anterior toothless portion of premaxillae 
short, usually shorter than length of tooth row. Nasals of moderate 
length, roughly as long as dental row, and wedged posteriorly into 



249 

182 frontals, almost at a right angle; anteriorly, each individual bone is 
deeply notched at the center, i.e., anterior end has a furcate appear- 
ance. Orbits relatively large. Lacrimals fairly large with rather large 
depression for preorbital gland, which occupies much of the frontal 
part of the bones even though very small. Tympanic bulla with an 
uneven surface and a short auditory meatus (its length invariably 
shorter than that of third upper molar). Anterior end of jugal not 
enlarged and does not reach lacrimal duct. 

Upper canines absent in most animals; small canine seen rarely 
on one or both sides in some males. 

In addition to marked age-related, sex-related, and seasonal vari- 
ations, the size and weight of roe deer also undergo considerable 
geographic variability. Females are invariably smaller and lighter 
than males, usually weighing about 16% less than males in a given 
locality. The geographic differences are so great that members of the 
largest subspecies weigh roughly 1.5 or even 2.0 times more than 
members of the smallest subspecies. Body length varies from 100 
to 155 cm, height at withers from 75 to 100 cm, maximum skull 
length from 180 to 253 mm, and weight from 20 to 59 kg. (V.H.) 

Taxonomy 

The genus contains only one species. For more details see 
characteristics of subfamily and genus. 

Geographic Distribution 

Genus Capreolus is distributed in western Europe, western part of 
Near East, southern half of European part of the USSR, southern 
Siberia, the Far East, Kazakhstan, Tien Shan mountain range, Mon- 
golia, northeast, east, and southeast China, and south to Szechwan. 

Geographic Range in the Soviet Union 

The range of roe deer in the Soviet Union is extremely extensive, 
covering all except the extreme west, southwest, and southeast parts. 
In the USSR the range is associated with the southern rim of the 
forest zone and forest-steppe, partly with the steppe zone, and mon- 



Differences in linear dimensions are invariably much less and not so obvious. 
Difference in weight of antlers is, perhaps, relatively even more than that of body 
weight, since antlers are relatively larger in large Siberian roe deer and in some other 
forms of this type. 



250 

tane forests. In its natural form the historic range was represented by 
one single zone although the dispersal of the animal at some places 
was sporadic or it deserted some places in winter or for a few years. 

The boundaries of some parts of the range of roe deer, primarily 
its northern boundary, show a marked dependence on climatic con- 
ditions. The northern boundary in particular is associated with 
depth of snow. It has been demonstrated that a mean maximum 
snow height of 50 cm restricted the dispersal of species in northern 
European and Asiatic portions of the USSR (Formozov, 1946). This 
association is, however, only of a general nature; there is no strict 
correlation of range boundaries with the foregoing isogram. At sev- 
eral places the zone of regular habitation of roe deer extends consid- 
erably beyond, into places with a snow depth of 60 or even 70 cm 
(western Pri-Urals and western Siberia). Sometimes a temporary 
shift in range beyond this line took place for several consecutive 
years due to various causes, for example, a series of less snowy win- 
ters and reduced interference by man (Nasimovich, 1956). Irregular 
visits of some individuals or groups of animals beyond the above- 
mentioned isogram into places where conditions of living are favor- 
able is a fairly regular phenomenon. Nevertheless, the importance of 
snow for demarcating the northern range boundary is fairly clear; 
significant nortfiward projections of boundary are associated with 
relatively less snowy zones (Pri-Baltics, Urals, Trans-Urals, Pri- 
Baikal, and others), and withdrawal southward in more snowy zones 
(central zones). 

During the historic period the range of roe deer underwent sig- 
183 nificant variations. Essentially these changes are similar to those 
seen in the case of wild boar described earlier. This is largely 
explained by the similarity of the northern range boundaries for 
both species and the causes of their fluctuations under the influence 
of the same set of factors (hunting by man and at times by predators) 
influencing identical peculiarities of the species in identical ecologi- 
cal conditions (response to snow). In many respects these differences 
are quite diverse both in nature and magnitude. 

The main range changes in roe deer in the period under discus- 
sion are as follows. The natural range prevailing evidently up to the 
seventeenth and early eighteenth centuries was continuous and 
extended from the western boundaries of the [Soviet] state through 
the basins of the Dnieper, Don, and Volga into the Urals, and farther 
into western Siberia up to the Pacific Ocean. In the south the range 
ran into the Crimea and the Caucasus. Later, in the nineteenth 
century, the range began to shrink, mostly in the European part of 
the Soviet Union and also in the Urals and western Siberia. This 



251 

process was considerably intensified in the latter half of the nine- 
teenth century. It proceeded particularly intensely in the twentieth 
century. The culmination of this process and a corresponding reduc- 
tion of range occurred in the 1920's. An essentially similar state 
prevailed by 1930, although a tendency toward expansion was 
observed at some places. 

By then, roe deer were absent in much of the European part of 
the USSR. They evidently disappeared from considerable areas to 
the east in western Siberia, in Kazakhstan, etc. Thus, at that time the 
range not only contracted but was also broken down into several 
zones which were totally or partly isolated from each other. The 
resultant zones could be called the western, Crimean, Caucasian, 
Ural (partly also west Siberian), Siberian-Far East, and Middle Asian 
locales. Division of the range into western, Crimean and Cauca- 
sian zones and separation of the Ural-Siberian zone occurred long 
ago, while isolation of the Ural and Siberian-Far East zones, and 
evidently that of the Middle Asian zone as well, occurred in the first 
two decades of the present century. 

Commencing from the 1930's an expansion of the range of roe 
deer has been observed, colonization of the animal at some places 
being quite rapid. The most marked changes occurred evidently in 
Yakutia, Pri-Urals, and western Siberia. The Ural region of habita- 
tion fused with the Siberian-Far East and the Middle Asian habitat 
also joined later. However, fusion of the western, Caucasian, Cri- 
mean, and Ural ranges has not occurred. 

Reconstructed range. The northern boundary of the recon- 
structed geographic range'^^ in the USSR ran northwest along the 
shore of the Gulf of Finland, encompassed the Karelian Isthmus, 
and extended into Finland. '''° Later it ran along the southern bank 
of Lake Ladoga roughly up to the mouth of the Syasa. From there it 
turned southeast and later east, running through Tikhvin, the 
Rybinsk reservoir, and Yaroslav (or slightly more northward, 
toward Danilov), to Kostroma, and the region slightly north of Gor'- 
kii (Balakhn), or close to Semenov. From there, for some inexplica- 
ble reason, it turned toward Kazan or slightly more north. 

Some of the more advanced peripheral points through which the 
boundary line extended represented the limits of transgressions of 

'"in the present case the term "reconstructed geographic range," as will be 
explained later, connotes a meaning different from the normal sense; the reference is 
more to the maximum range over the historic period. 

'''°In the Karelia Isthmus roe deer were sighted rarely and occurred there only as a 
visitor. The boundary of the more regular habitat ran along Neve. 



252 

the animal or the maximum northward advancement of the bound- 
ary in several years with poor snowfall. This pertains primarily to 
deer occurrence in Yaroslav and Kostroma districts and in the region 
between Kostroma and Gorkii. Nevertheless, there is no doubt that 
the boundary of regular habitation of the animal in the Yaroslav- 
184 Gorkii section also came quite close to the above line; roe deer 
colonized fairly extensively in the Moscow district even in the six- 
teenth century, and still survived in Podol and Serpukhov districts 
and even the Bogodor district in the latter half of the last century, 
and were seen around Moscow itself. Sightings are also known from 
around Pereslav-Zalessk; roe deer existed normally and in considera- 
ble numbers even quite recently in the Gorbatov region, southwest 
of the Oka mouth, on its right bank. They resided permanently 
north of Tula (Tula Zaseka) and in regions adjoining the Volga 
from the south, in the zone between the Oka mouth and Kazan 
(southern part of Nizhegorod and Simbir provinces). 

From Kazan the boundary ran east roughly along 56° N lat. or 
along the Kama, emerged toward the mouth of the Beloe into the 
Kama, and departed beyond the Kama. From there it turned steeply 
straight north and ran along the PriTIrals through Регт,"*' 
Dobryanka (about 80 km north of Perm), Kizel, and Cherdyn' into 
the sources of the Pechora, up to the discharge of the Ilich into the 
Pechora (62°30'). Nevertheless, the region of the source of Pechora 
evidently is only a region of occasional visits by the animal. 

In the Urals per se the northern boundary of the range fell on the 
latitude 62° N lat., evidently even slightly more north at 62°30' N 
lat. Sightings have been reported in the eastern foothills up to 64° N 
lat. (upper course of the Lyapin, a tributary of the Severnaya Sosva). 
It is possible that animal visitations may have extended even farther, 
but there is no precise information about this. These places repre- 
sent some of the northernmost points of occurrence of roe deer.'"*" 

In the Trans-Urals the range boundary turned sharply south 



The position of the range boundary between Menzelinsk, the Ufa River, and 
Perm is not clear. Possibly, it proceeded roughly along the Kama. 

'"^The range boundary in the European part of the USSR and the Urals is based 
on data from M. Bogdanov, 1871; Lavrov, 1929; Bobrinskii, 1944; Middendorff, 1869; 
Chernaya, 1860; Kessler, 1850; Formozov, 1946 and 1947; Severtsov, 1955; Ognev and 
Vorob'ev, 1924; Danilov, 1947; Nasimovich, 1956; Fedesov and Nikitin, 1951; Kuklin, 
1951; Kirikov, 1952, 1953, and 1955; Heptner. Morozova-Turova and Tsalkin, 1950; 
Korneev, 1952; Serzhanin, 1955; Elpat'evskii, Larina and Golikov, 1950; Schwarts, 
Pavlinin and Danilov, 1951; Popov etal., 1954; Flerov, 1952; Eversmann, 1850; Pallas, 
1811; Demidov, 1940; Turkin and Satunin, 1900; V.N. Skalon, 1946; Siivonen, 1953; 
Kalela, 1948; and others. 



253 

from the upper reaches of the Lyapin in the upper reaches of the 
Severnaya Sosva, and from there to the east or southeast to the Ob', 
where it merged roughly at 61°40' or 62° N lat. (Leushi), i.e., 
markedly north of the mouth of the Irtysh. From there the range ran 
south along the left bank of the Irtysh up to a point slightly south of 
the confluence of the Tobol with the Irtysh. Later it turned sharply 
east, intersecting the Ishim slightly above its mouth and the Irtysh 
below Тага. After passing through the northern parts of the Bara- 
binsk steppe, it intersected the Ob' and Tom' slightly south of the 
latter. Roughly at the same latitude the boundary ran toward Kras- 
nodar, but slightly short of reaching it rose up to 58° N lat. on the 
left bank of the Yenisey (regions of Bol'shaya Murta, Kazachinsk, 
and Pirov). Crossing over at this level onto the right bank of the 
Yenisey, the boundary turned east along the Angara, intersected it 
somewhere below the Ilim mouth and ran farther through the 
Katanga River (at the source of the Podkamennaya Tunguska to the 
northeast of the Ilim mouth), roughly around 59° 30', and through 
the source of the Nepa (tributary of the Nizhnaya Tunguska at its 
source). 

From there, in a manner not known for certain, but evidently 
following the Lena, the boundary turned northeast and crossed onto 
the left bank of the Lena, above (west of) Mukhtuya roughly at 1 13 
to 114° E lat.'"*^ Later it ran into the Vilyui in the region of western 
Suntar. Thus, in this part of Yakutia, roe deer lived in the basin of 
185 the lower reaches of the left hand tributary of the Lena and Nyuya, 
most probably in much of the basin, if not the whole of it. On 
intersecting the Vilyui the boundary encompassed the left bank of 
the river within the Suntar bend and ran roughly toward the mouth 
of the Markha. Farther away in the east it turned generally along the 
Vilyui possibly also encompassing the lower reaches of its left hand 
tributaries, and emerged into the Lena in the region of the mouth of 
the Vilyui, i.e., around 64°, or slightly more north.''*'* It is possible 

Abundant information is available for some parts of the range of roe deer in the 
zone between the Yenisey and Baikal and for western, northern, and northeastern 
Pri-Baikal; it exceeds the information available for several regions in the western 
parts of the Soviet Union. However, much of the information is contradictory. 
Because of complex orography, different local conditions of snowiness, and so on, the 
range there can be regarded as extremely complex and, what is more, gaps are evident 
in this information. Moreover, changes there could vary quite intensely in different 
years. Information is absent or scanty for vast expanses, for example the regions of the 
Kirenga, Chai, Chui, and Mama basins, and elsewhere. 

In Yakutia roe deer occupied the "Vilyui" region, Lensk, Olekminsk, Ordzhonikidzev, 
Gornyi, and Yakutsk regions (Belik, 1953; and other sources). According to other 
authors (D.U. Ivanov) roe deer did not go beyond the Vilyui in the north and did not 
reach even up to its mouth (known as Kobyai or Kobyaitsa). 



254 

that there the boundary ran even more northward since encounters 
with roe deer are known in the area of Syalakh (Sylakh) in the 
Zhigansh region at 66° N lat. This is the northernmost recorded 
habitat of roe deer in the USSR. 

On the right bank of the Lena roe deer occupied nearly the whole 
area between the Lena and Amga, in the north up to the latitude of 
the Amga mouth into Aldan, and in the east and south up to, or 
almost up to the Amga, from the mouth up to the source. From the 
source of the Amga the southern boundary on the right bank of the 
Lena turned southwest and crossed straight to the Olekma. Thus, in 
Yakutia the range of roe deer was like a large promentory to the 
north. Such a distant penetration of roe deer into northern Yakutia, 
apart from the reasons stated above (snow), was associated and lar- 
gely coincided with the entire complex of forest-steppe and steppe 
topographies developed in this part of Yakutia (along the Vilyui). 

Upon crossing the Olekma the boundary along this river turned 
south toward the western extremity of the Stanovoy range. From 
there it turned sharply east and, for some inexplicable reason, 
extended toward the Amur basin'''^ evidendy along the northern 
slopes and foothills of the Stanovoy range through the uppermost 
reaches of the Aldan (Timptonsk region of Yakutia with the center 
at Chul'man), the watershed of the Gonam and Algama, and the 
upper reaches of the Uchur. The range, however, did not cover the 
Aldan-Uchur range (K.A. Vorob'ev). 

In the east the boundary did not reach the Pacific Ocean at these 
latitudes, but on intersecting the Stanovoy range or the western 
extremity of the Dzhugdzhur turned southeast and south, encom- 
passing the upper reaches of the Uda and running along its right 
hand tributary, the Galam (roughly at 54°30' N lat. and 133°30' E 
long.; V.H.). From there the boundary line excluded the lowermost 
reaches and mouth of the Amur outside the range, and ran into the 
mouth of the Amgun on the Amur, and from there evidendy 
through the region of Lake Kiza toward Tatarskiy Strait, south of De 



""'There is not much accurate information for northern Yakutia where roe deer 
became estabhshed in the last decade. Nevertheless, the data and map given here 
represent significantly more accurately the actual picture of the range north and east 
of Lena than those published earlier (Lavrov, 1929; Bobrinskii, 1944; Flerov, 1953). 

'■'^Information given for the Aldan basin is based on reports which are partly 
inaccurate and in which geographic names have obviously been distorted. Thus, it 
has been shown (V.N. Skalon, 1937) that roe deer were caught on Meguskaya River 
and also on the watershed of the Gonata and Algata. Middendorff (1869) reports that 
roe deer were found only on the southern slope of the Stanovoy range. 



255 

Kastri Bay, roughly at 51°. Sakhalin did not fall within the range. '"^ 
The southern boundary of the reconstructed range of roe deer in 
the European part of the USSR evidently traversed the Black and 
Azov Sea coasts. Although direct references to this are not many, 
based on the sum total of information about the nature of the Soviet 
south in the past and the ecology of roe deer, there is every reason to 
believe that roe deer occupied not only the forest-steppe, but also the 
whole of the steppe zone. The distribution in the latter was, of 
course, sporadic even as it is now in Kazakhstan (see below). There is 
187 no doubt that roe deer performed even longer migrations through 
the steppe (covering several dozens and hundreds of kilometers). 
This part of the range crossed directly into the Crimea and joined 
with the Caucasian part; in the south, in the Trans-Caucasian 
region, the range ran beyond the state boundary and also obviously 
covered the steppe region of eastern Trans-Caucasus. 

In the zone falling between the Don, Altai, and Tien Shan, i.e., 
in the region adjoining the northern part of the Caspian Sea, in 
Kazakhstan and Middle Asia, the southern boundary of the natural 
range was fairly complex and its features are still not wholly under- 
stood. In general the range there encompassed not only the steppe 
but also the rim of the semidesert, or even the desert zone. It is quite 
natural that the dispersal of the species there also remained sporadic. 
The mountain systems mentioned before also fell within the range. 
Adequate positive information is not available about the disper- 
sal of roe deer in the past in the Pri-Caspian region and any bound- 
ary drawn would only be approximate. Commencing in the west 
on the coast of the Caspian Sea in the region of the estuaries of 
Kuma and lower Manych, the range boundary evidently passed 
along the Don or its left hand tributaries, bypassed from the west the 
semideserts on the right bank of the lower reaches of the Volga, and 
ran toward the Volga in the Stalingrad region. It is possible that roe 
deer descended right up to the sea along the Volga valley in the 
distant past. In some manner not yet clearly understood the bound- 
ary turned east from the Volga and evidently covered the Eruslan 
basin and the region of the upper and middle Uzen'. It is highly 

The northern boundary of the range of roe deer in the Asiatic part of the Soviet 
Union has been taken from Nasimovich, 1955; Yanushevich and Blagoveshchenskii, 
1952; Slovtsov, 1892; Zverev, 1937; Kopylov, 1948 and 1950; Favorskii, 1936; Podarev- 
skii, 1936; Abramov, 1954; Formozov, 1946; Tugaiinov, Smirnov and Ivanov, 1934; 
Troitskii, 1930; Novikov, 1937; Yakovlev, 1930; Gassovskii, 1927; Middendorff, 1869; 
Skalon, 1937 and 1946; Turkin and Satunin, 1900; Schrenk, 1859; Flerov, 1935 and 
1952; Maak, 1859; Abramov, 1954; Belyk, 1953; Fetisov, 1953; Lavrov, 1929; and some 
others (including the unpublished material of D.U. Ivanov and A.A. Sludskii). 



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257 

probable however that along these rivers it descended even more 
southward to the Kamysh-Samar lakes. From there the boundary ran 
along the Ural [River] to Uralsk. Evidently, as along the Volga, roe 
deer descended far south along the Ural valley also. 

Farther away in the east the range covered a considerable part of 
Kazakhstan where roe deer were distributed (partly surviving even 
now) extremely sporadically, but extensively. Commencing at 
Uralsk the southern boundary of the range ran southeast, covering 
from the south the Ilek, including the Khobda basin, a tributary of 
the Ilek (Khobdinsk, Aktyubinsk region), and later left at Dzhurun 
(about 100 km south of Aktyubinsk). From there it turned almost 
straight east, passing roughly between the towns Irgiz and Turgay 
and later, slightly descending southward, encompassed the Ulutau 
mountains from the south. 

Traversing through the upper part of the Sarys River and later 
through the regions of the Aktau and Ortau hills along the northern 
coast of Lake Balkhash at a distance of about 100 km from it, 
roughly at the meridian of the eastern extremity of the lake, the 
boundary turned south and ran toward the lake. The range left out 
not only the Chingiztau and Tarbagatai but also the whole of the 



'"^In the steppes roe deer occupy not only large forest masses but also small groups 
of isolated forests, ravines, areas with shrub thickets in the plains as well as in hilly 
terrains, forests in river valleys, sometimes overgrown river banks, and reed thickets of 
lakes. They often live in very small sections even when separated from each other by 
several dozens and even hundreds of kilometers of open steppe (pine forests of Ara- 
Karagay, Aman-Karagay, Naurzum-Karagay, forest sections around Aktyubinsk, Ulu- 
tau mountains, Kokchetav, Kent mountains, Kuu. Karkaralinsk, Airtau, and others). 

'""The information that in the Turgay basin region roe deer were confined to and 
hunted in the southeastern part, i.e., east and northeast of the Aral Sea is clearly 
incorrect (Turkin and Satunin, 1900). Turkin in Zver'e Rossii (Animals of Russia) 
used information about fauna and hunting submitted by the provincial governments 
to the Commission for preparing hunting legislation, of which Turkin was a member 
in the 1890's. The provincial governments, however, did not always examine well the 
data given them. In the present case some confusion with saiga seems indicated. 



186 Fig. 61 Geographic range of roe deer, Capreolus capreolus L. in the USSR and some 
of its modifications (scale in km). 

1 — boundary of reconstructed range and its maximum expansion in the last decade 
(mid-1950's); 2 — boundary in mid-1950's where it does not coincide with reconstructed 
range; 3 — boundary during its maximum reduction in 1920's (roughly by 1930) in the 
European part of the USSR, Urals, and Siberia; 4 — places of roe deer visits toward 
Syalakh at 66°N lat. (south of Zhigansk on the Lena). Additional details given in Figs. 
62 and 63. V.G. Heptner. 



258 

Alakul'sk basin, and crossed the Dzhungarsk Alatau into the Central 
Asiatic part of the range. 

Thus in Kazakhstan roe deer occupied not only the entire steppe 
zone in general, but also very significant portions of the semidesert 
zones, penetrating even into the deserts. As mentioned before, in 
these zones they took shelter in some of the apparently least suitable 
areas. Elevated sections with trees or at least shrubs were the most 
favorable. It is interesting that in Kazakhstan roe deer penetrated far 
into the regions of dispersal of such typical steppe, semidesert, and 
desert ungulates as saiga, goitered gazelle, kulan [wild ass], and wild 

,150 

horse. 

The range of roe deer covered a considerable region in the moun- 
tains of Middle Asia in the east. The boundaries of this zone are, 
however, as yet poorly understood and can only be generalized. A 
fairly accurate picture can be drawn only in the north. There, in 
spite of its fairly complex features, it encompassed the Dzhungarsk 
Alatau, intersected the Iliisk basin, ran into the Trans-Iliisk Alatau, 
and later into the Kirgizian mountain system. Along the Karatal and 
Hi Rivers the range formed narrow projections right up to Balkhash 
and along the Chu up to the floodplains of its estuary zone. The 
Chu-Iliisk mountains in the north, including the Dzhambul moun- 
tain group, fell within the range. Farther away the boundary made a 
prominent projection northwest, encompassing the Karatau moun- 
tains. From there, in a manner not precisely understood but 
obviously complex, it proceeded south, bounded on the west by the 
mountain ranges of the Tien Shan system, later ran along the north- 
ern slopes of the Ferghana valley, turned east, and evidently ran 
toward the state boundary with China, exiting from Soviet frontiers. 
The southern boundary of the range of roe deer in the mountains of 
Middle Asia evidently proceeded in the same manner.'^' Informa- 
tion is available that the habitat of this species extended more south- 
erly, in the Gissar mountains (Flerov, 1935; Turkin and Satunin, 
1900) and even the Darvaz range (Lavrov, 1929). These statements, 
sometimes with additional data, are based on information provided 



'^°The range boundary of roe deer in the section under description is based on data 
from V.G. Heptner; Kuznetsov, 1948; E.I. Strautman, 1953; Sedel'nikov, 1903; 
Antipin, 1941; Lavrov, 1929; Turkin and Satunin, 1900; Derevyagin, 1930; Lentovs- 
kii, 1928; Korsakov, 1938; Selevin, 1925; N.V. Pavlov, 1931; and others (mainly the 
unpublished data of A.A. Sludskii). 

'^'References to the northern part of the eastern Bukhara basin, Trans-Caspian 
region, and Tien Shan south of the Pamir (Flerov, 1935) have been omitted in the 
present study. 



259 

by G.E. Grum-Grzhimailo (1886, pp. 92 and 93). The first to use this 
information was Turkin, who misinterpreted it (Turkin and 
Satunin, 1900). Like Turkin, other researchers also erred because 
though they referred to the original source they evidently used Tur- 
kin's text. Yet from the original data of Grum-Grzhimailo it is 
absolutely clear that what he had in mind were wild goats (Capra — 
evidently С sibirica and С falconeri)}^^ The assertion that "roe deer 
inhabit the Zeravshan mountains" (Flerov, 1935) has not only not 
been confirmed but is contradicted by new data on the total absence 
of roe deer within Tadzhikistan (V.I. Chernyshev). 
189 Along the Syr-Darya, at least in the period under consideration, 
roe deer were evidently absent and the suggestion of their occurrence 
in the valley above Kzyl-Orda (Antipin, 1941; map) is unfounded. In 
the very distant past it may perhaps have been sighted on the river 
but there are no references to this possibility. Information about the 
occurrence of roe deer in the Amu-Darya estuary (Lavrov, 1929; 
according to report of Manteifel') is incorrect. Statements that in 
Samarkand district roe deer thrive in abundance and "In the Trans- 
Caspian district sizable numbers lived in the Krasnovodsk region" 
(Turkin and Satunin, 1902) are erroneous and evidently the result of 
confusing mountain goat and goitered gazelle (see footnote 152). 
Also erroneous, in part simply fantastic is the suggestion that roe 
deer inhabited the Bol'shoi Balkhan early in the last century and 
were sighted even in 1941 and 1942 (Bazhanov, 1945). 

Throughout the stretch from Semirech'ya to the Pacific Ocean 



Lavrov (1929) states that Grum-Grzhimailo in 1886 pointed out the presence of 
the deer in Gissar(! V.H.)and Darvaz mountains, and that residents in those days were 
well acquainted with it. Grum-Grzhimailo's text states: "In the Gissar mountains 
occur two species of goats, but no kind of deer. There are also none in the Darvaz and 
nearby, where the residents are acquainted only with the kiik, consequently, once 
more in them there are only mountain gOats" (p. 92) "Capra sp.? — kiik in Kirgiz" 
(p. 93). 

Yet it is quite possible that roe deer inhabited or could have occurred in the 
Alaiskii mountain range. In the zoological museum of the Academy of Sciences there 
is a skull of a specimen caught by Grum-Grzhimailo on the northern slope of the 
Alaiskii range on Dzhusala River in 1889. It has not been possible to establish pre- 
cisely the place at which the animal was caught. It is highly probable that the place of 
capture was the eastern or northeastern extremity of the mountain range where it 
adjoins Ferghana, i.e., in the region of the sources of the Kara-Darya or in the 
southern part of the mountains which surround the Ferghana valley from the east. 
This does not alter the conclusion drawn above about the southern range boundary of 
roe deer in the mountains of Middle Asia. In the same part of the mountains which 
enclose Ferghana valley from the south roe deer were evidently absent, at least in the 
region south of the city of Ferghana (V.G. Heptner). 



260 

the range of roe deer extended up to the state boundary in the 
south. 

Geographic range in the period of utmost depletion. As shown 
above, in this century, and in particular in the nineteenth, due 
directly to the extermination of animals over considerable regions 
the range underw^ent steady shrinkage. Separation of the Caucasian 
and Crimean parts of the range from the European occurred long 
ago. It took place notably in the latter half of the last century and the 
early part of the present century. The maximum reduction of the 
area of the range occurred in the 1920's. By this time the range w^as 
already divided into several smaller parts, as mentioned before, and 
their outlines in most cases cannot be determined precisely. 

The eastern boundary of the western habitat of roe deer com- 
menced at the eastern extremity of the Gulf of Finland and formed 
an irregular and fairly complex line before turning south and leav- 
ing the USSR to enter Rumania. From Leningrad the boundary 
proceeded east along the Neva and the southern bank of Lake Lad- 
oga roughly up to the mouth of the Syasa. Visits into the Karelian 
Isthmus did take place but very rarely. From the Syasa estuary this 
line, without crossing the river in the east, turned south and ran 
toward Borovichi, later slightly southeast toward Vyshnemy Volo- 
chek, and from there straight south into Ostashkov. Later it dipped 
slightly southwest into Velikii Luki from where it turned south- 
southeast toward Smolensk and Zhizdr, and later followed a fairly 
steep westward arc, running initially southwest into the Novozyb- 
kov region, and later still turned steeply eastward toward L'gov, and 
south-southeast of it into Graivoron, more to the west of Belgorod. 

From there the long penetration of the range east and southeast 
into the Kharkov region commenced. From Graivoronsk the bound- 
ary line proceeded southeast of Valuika, bypassing Kharkov from 
the north, turned to Chuguev, and later passed into the Izyum 
region. From there it turned sharply northwest and proceeded to 
Zmiev and Krasnokutsk (west of Kharkov, south of Akhtyrka), and 
later into the region south of Konotop. From there the boundary 
turned west, passed south of Nezhin and, taking a steep southwest- 
ern or even southern direction, proceeded toward the Dnieper south 
of Kiev. It traversed some distance along the river and intersected it 
evidently at about 50° N lat. 

On the right bank the boundary ran almost straight south into 

The range in the Tien Shan and Pamir-Alaiskii systems is based on data from 
E.I. Strautman, 1953; Kuznetsov, 1948 and 1948a; Antipin, 1941; Turkin andSatunin, 
1900; Grum-Grzhimailo, 1886; Flerov, 1935 and 1952; Lavrov, 1929; and other sour- 
ces, including original data from V.G. Heptner and V.I. Chernyshev. 



261 

the region south of Korsun'-Shevchenkovsk. It then ran farther 
southwest, south of Uman, intersected the Bug somewhere in the 
region of Pervomaisk and ran into Moldavia, evidently passing there 
in the region of Kishinev and later into Rumania. Outside the fore- 
going boundary lay a very large pocket of habitation of roe deer 
north of Voronezh in Usmansk forest (Voronezh preserve). Further- 
190 more, there were evidently some small pockets of habitation of roe 
deer on the right [west] fringe of the Ukraine, for example in the 
Black Forest of Kirovograd district, and on the left at Novomoskovsk 
near Dnepropetrovsk. The occurrence of deer at Izyum was also in 
the form of an isolated pocket. 

The above boundary line has been described only schemati- 
cally.'^^ Information on the range of roe deer is not only scanty, but 
also for the period in question the contraction of the boundary was 
very indeterminate. All along the eastern edge of the range, over a 
goodly portion if not all of it, roe deer populations were sporadic 
and small pockets of these animals often widely separated. The 
number of animals in these pockets, since the total population of the 
species was far from large, was understandably quite small. The 
boundary of this zone varied notably in different years due to animal 
extermination at some places and range expansion in others. At 
places colonization commenced evidently even during the 1920's but 
was observed somewhat later. Finally, there are records of visitations 
east of the above line, often over fairly considerable distances. In the 
west the boundary of this region extended up to the Baltic Sea and 
into the Carpathians and Rumania. 

Distribution in the Crimean region was restricted to a small area 
in the forest section of montane Crimea. 

Distribution in the Caucasian region during the period under 
consideration was roughly the same as in the 1950's (see below). 

Distribution in the Ural-western Siberia was fairly complex. The 
westernmost point lay in the region slightly east of Ufa. From there 
the boundary proceeded north and northeast, generally along the 
Ufa River or parallel to it, but barely west of it. From Krasnoufimsk 
it turned northeast and ran into the upper reaches of the Chusovoi 
and ascended along the Urals roughly up to 58° or slightly more 
northward. It then arced southward, turning southeast and later 
east, through Irbit, north of Tyumen, and south of Tobol. Running 
in that direction and slightly dipping south, it reached the vicinity 
72° E long. From there, forming a promentory jutting into the east, 

' Mainly according tadata of Lavrov (1929), which provide additional details not 
given here. 



\0 'S 20 25 30 35 40 45 50 55 60 65 70 75 




200 200 400 600 k 

''■■'' I I 1 



"" О 5 



'-°" 3 ^ 6 



263 

the boundary turned steeply southwest. Passing through the city of 
Ishim it proceeded in the form of an irregular line in the same 
direction into the region south of Kurgan. 

The southern boundary of this region of habitation of roe deer 
rose steeply north from the southern extremity of the Urals roughly 
up to the latitude of Troitsk on the eastern side of the forest zone of 
the range. From there it turned sharply east, passed through Troitsk, 
intersected Tobol, formed a projection southward along the right 
bank of the Tobol up to the latitude of Kustanai, and a similar 
projection from the region south of Kurgan to Kokchetav. 

Around this period of extreme contraction, a break in the range 
occurred along 55° N lat. for a distance of about 1 ,500 km or roughly 
24° between the western point of distribution in the Urals and the 
eastern boundary of the western region (excluding the narrow pro- 
jection in the Kharkov region and the section at Voronezh), which 
formerly represented a continuous range. The Ural-western Siberia 
distribution by then was also separated from the range covering the 
rest of Siberia; in the region of Omsk, Novosibirsk, and the Pri-Altai 
steppes roe deer likewise disappeared. The interruption of the range 
there was somewhat less than in the European part of the USSJR.. 
Precise data are few and while it is possible that the interruption of 



''* The boundary of Ural-western Siberia distribution is based mainly on data 
from Lavrov (1928), while that for Troitsk, Kustanai, Kurgan, and Kokchetav is based 
on unpublished data compiled by V.G. Heptner. 

The outlines of indi\'idual occurrences given here and at some other places later 
are somewhat schematic and generalized. Some details are available in Lavrov's work 
(1929) and in faunistic studies of the 1920's and early 1930's. 



191 Fig. 62. Geographic range of roe deer, Capreolus capreolus L. and its changes 
in the European part of the USSR and the Caucasus (scale in km). 

1 — boundary during utmost reduction of range by 1930 (mostly in the first half and 
middle, and partly at the end of the 1920's; this boundary in the Crimea and the 
Caucasus corresponds also to the boundary depicted in no. 3); 2 — isolated pockets of 
occurrence of roe deer in the 1920's — Voronezh sanctuary, Novomoskovsk region, and 
Black Forest; 3 — boundary of range at the time of its maximum enlargement in the 
mid-1950's; 4 — northern and southern boundaries of the reconstructed range of roe 
deer where it does not coincide with the boundary for the mid-1950's; 5 — some distant 
transgressions of roe deer after the 1930's— Balakhna, the mouth of the Kama, Volsk 
and Karelia Isthmus; б — region of occurrence of roe deer of unidentified origin 
(partly acclimatized) in northern Moscow region. Question marks indicate probable 
habitats in the past (reconstructed range) along the Ural valley (same for Volga). V.G. 

Heptner. 



264 

192 the range in the west of the European part of the USSR and Crimean 
and Caucasian sections represents the resuh of processes occurring 
over a very long period, the eastward interruption in western Siberia 
arose quite recendy, just before and during the 1920's. 

The Siberian-Far East distribution of roe deer commenced in the 
west in the Akai. The range boundary encompassed this area in the 
form of an arc commencing at the boundary with China. It traversed 
somewhat north of Zaisan, proceeded northwest sUghtly away from 
Irtysh, and then turned northeast in the form of an arc somewhere 
east of Semipalatinsk. Passing south of Biisk it turned into Stahnsk 
and later into Krasnoyarsk. In the Krasonyarsk region the northern 
boundary did not cross the Sibirsk [Trans-Siberian] railway line. 
East of the Yenisey it proceeded roughly at the same latitude 
through Aginsk region, slightly north of Taishet, and left at the 
Angara, roughly at Bratsk. Farther away the boundary turned 
markedly southward and intersected the Lena at Zhigalov as well as 
the Kirenga roughly at the same latitude. Turning northeast from 
there the boundary line extended somewhat north of the northern 
extremity of Baikal. Inclining somewhat south it encompassed the 
southern part of the North Baikal highlands and the upper reaches 
of the rivers of the Patom highlands, intersected the Vitim consider- 
ably south of Bodaybo,'^^ and ran toward the middle reaches of the 
Olekma. From there it turned south toward the Stanovoi range. The 
details of its position farther east are not clear. Evidently it pro- 
ceeded along the southern slopes of the mountains, but it is possible 
that it lay slightly more to the north as described above (see the 
outline of reconstructed geographic range). In any case there are no 
significant differences. 

The Middle Asiatic portion of the range in the period of utmost 
depletion differed from the one described above in that roe deer were 
evidently absent in the Zaisan depression and Alakul'sk basin; this 
part of the range was thus isolated from the Siberian-Far East part. 
Roe deer evidently did not go that far down along the Balkhash river 
system valleys (Hi and Karatal) and probably did not reach up to 
Balkhash. The animals did not penetrate the Chu-Ili mountains, at 
least they did not thrive that far north, nor did they emerge far into 
the plains along the Chu River. At Karatau also they evidently did 
not penetrate that far north. 

Geographic range in the 1950's. The period of extreme depletion 
of population and reduction of the range commenced, as mentioned 

'" The boundary up to the Yenisey is taken from the data of Lavrov (1929), and in 
Trans- Yenisey Siberia from Favorskii (1836). 



265 

above, during the 1920's, more precisely from the first half of that 
decade. The position was essentially similar by 1930 and in the early 
years of that decade. Some efforts toward expansion and restoration 
of the range were begun in the 1920's (in the middle and latter half of 
that decade); measures for conservation were initiated here and there 
but proved ineffective in altering the overall pattern of depletion. 
This is true primarily of the European part of the country, the Urals, 
and western Siberia. 

Range expansion, leading to the present situation, became 
detectable in the 1930's. This process developed better in the 1940's. 
and continued intensely in the 1950's. The principle of expansion in 
various parts of the range is far from similar, however. In the Euro- 
pean part of the USSR colonization in the western section has and is 
taking place in the east, partly in the south, in regions where the 
animal was formerly present, i.e., there is only a restoration of the 
range. It is proceeding extensively and quite intensely over a large 
territory, but is still far from realizing the former maximum range. 
The boundary has been restored only in a small section in the north- 
west. It has not been possible to identify the influence of any natural 
stimuli for colonization. The more important factors are undoubt- 
edly lessened persecution by man and attempts at animal conserva- 
tion (reacclimatization is not taken into consideration here; for this 
aspect see below). 

In the Caucasian and Crimean parts of the range there are practi- 
cally no settlements at all, and the position of the range in the last 
decade, to tell the truth, has not altered. 

193 Colonization in the Urals, western Siberia, and Kazakhstan is 
proceeding intensely in all directions (see below). This also means 
the restoration of the former range, which has already reached its 
former maximum limits. In some parts there is even an enlargement 
of the range compared to its former boundaries. This process, occur- 
ring under the direct or indirect influence of man, is also associated 
with natural factors. One primary factor is the warming of the cli- 
mate over a vast area. Thus, two processes have taken place in the 
past decade in the Urals and western Siberia and the outlines of the 
range given above for the zone under consideration represent more 
of the maximum than the reconstructed range (the latter term usu- 
ally designates the maximum extent of the range in the past). As a 
result of colonization, mainly from the west, distribution in Ural- 
western Siberia has merged with the Siberian-Far East range. 

194 At present in central Baikal, although some enlargement of the 
range has been observed, the former northern limit has not yet been 
reached. In Yakutia in the last decade and the las.t few years intense 



266 





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70 










100 

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100 200 300 km 

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193 Fig. 63. Colonization of roe deer northward in heavy snow regions of the Urals 
and western Siberia (scale in km). 

1 — northern boundary of distribution in 1905 to 1915; 
2 — same in 1940 and 1941. Crosses represent northernmost points of transgres- 
sion in the last decade (up to commencement of 1950's). Broken line indicates 
long-term mean height of snow cover in cm (from Nasimovich, 1955). 



colonization has occurred and the range there now is much larger 
than the original range in the past, not only about a century ago but 
even in the 1930's and 1940's. It is in relation to this territory that 
there is utmost justification in referring to the range as maximum 
and not as reconstructed. The northward movement of roe deer there 
in the last decade was evidently the result of primarily natural 
causes. Farther away, in the east, i.e., in the region of the Stanovoi 
range and the Amur and Ussuri basins, colonization of roe deer has 
not occurred to any noticeable extent. 



267 

In the Middle Asiatic part of the range the process of enlarge- 
ment has taken place but on a small scale in terms of area. Neverthe- 
less, as a result of colonization from both sides, the Middle Asiatic 
and Siberian-Far East parts of the range merged. On the whole, the 
present range of roe deer is in a fluid state and a general tendency 
toward enlargement clearly manifested. However, the picture of 
range dynamics is not only complex but also different in different 
parts of the range. It is not determined everywhere by the same 
factors. In some parts these factors are purely man-made and in 
others natural or mostly natural, or a combination of both factors. 

As a result of the phenomena described for the last decade the 
range has not only increased in size, but some restoration of its 
continuity achieved. By the middle of the 1950's it was broken down 
not into six but only into four parts: western, Crimean, Caucasian, 
and Siberian-Middle Asiatic, in which the Ural, Siberian-Far East 
and Middle Asiatic parts had merged. If this tendency persists, the 
merger of western and Siberian-Middle Asiatic regions can be 
expected in the future. 

The range of roe deer by the mid- 1 950 's can be described as 
follows. The eastern boundary of the western habitat, commencing 
on the southern bank of the Gulf of Finland encompassed Karelia 
Isthmus, going from there into Finland, proceeding farther along 
the southern bank of Lake Ladoga, and reaching roughly the mouth 
of the Syasa. From there the boundary line ran southeast into Tikh- 
vin, then in the same direction into Rybinsk reservoir (Darvinov 
preserve), and into Yaroslav. In the section from the Gulf of Finland 
up to Yaroslav the range boundary has already been restored. Evi- 
dently associated with this part of the range are long excursions of 
roe deer as far as Balakhna northwest of Gorkii. 

From Yaroslav the range boundary turned steeply southwest 
toward Pereslav-Zalessk and from there westward into the Zavidov 
region (on the southern bank of the Volga reservoir intersected by 
the line joining Moscow-Leningrad). From here the boundary 



Roe deer have been encountered, though rarely, throughout northern, north- 
western, and partly western parts of the Moscow district, in the regions of Konstantinov 
(north of Zagorsk), Dmitrov, Klin, Vysokov (west of Klin), Volokolam, Novopetrovsk 
(east of Volokolamsk) (Danilov, 1947) and Shakhov (west and southwest of Voloko- 
lamsk; L.P. Rasshivalina), and probably in some other regions. 

It is possible that, at least partly, this was a part of the natural range formed as a 
result of colonization from the west and northwest. At the same time, however, there 
is no doubt that the Moscow portion of the range was formed to some extent by 
Siberian roe deer released time and again for purposes of acclimatization in the 
Zavidov region (those released in Pogonno-Losin Island in Moscow evidently per- 



268 

line turned south, passing somewhere along the area between the 
195 Moscow and Smolensk districts, possibly through northwestern 
region of Moscow district: Lotoshin, Shakhov, and Volokolam. 
Forming a broad arc it proceeded farther, evidently through the 
region west of Kaluga, Orel, or slightly west of it, and turned south- 
east to pass slighdy north of Voronezh (Voronezh preserve) and, 
at the mouth of the Voronezh River, into the Don.'^^ From there, the 
boundary line, for reasons not yet clear, proceeded east into the 
Khoper and encompassed the forest stretches in the river valley 
between Novokhopersk and Borisoglebsk and the lower reaches of 
the Vorona (Borisogleb forest, Tellermanov wood, and Khoper pre- 
serve). It is also possible that this section of the range was isolated. 
Visitations of roe deer, evidently from the west, were observed dur- 
ing the war years even more to the east, in the region of Vol'sk. 
From the Khoper, also in an inexplicable manner, the boundary 
proceeded southwest to Valuika and from there turned steeply south- 
east along the Derkul River, which flows into the Donets from the 
left near Lugansk, and proceeded toward a point slightly north of 
Belovod. Along the Derkul, the boundary ran up to its mouth where 
roe deer lived in Krasnoborsk forest. From there the boundary pro- 
ceeded northwest into the Izyum region and slightly inclining 
southwest turned into the Novomoskovsk (northeast of Dneprope- 
trovsk), intersected the Dnieper, and evidently ran through the Kiro- 
vograd district to the Pervomai and Kotov regions, then changed its 
westerly direction southwest and entered Moldavia. From Kotov it 
ran southwest to the Dnestr, descended along it to the south through 



ished totally), i.e., formed artificially. It is, however, impossible to establish this at 
present since the taxonomy of Moscow roe deer has not been studied. There is only 
one partially identified reference that in Zavidov European roe deer coming inde- 
pendently from the west were also found along with imported Siberian roe deer 
(Danilov, 1947). The presence of European deer could also be expected in the Voloko- 
lam region and Shakhov, as also in Klin and Vysokov. The dubious occurrence of roe 
deer in northern Moscow region has been depicted by a hatched line in the map. 

Information about the range of roe deer from Voronezh up to the border of 
Moldavia is scanty and very generalized. 

These data pertain to the 1930's (Izmailov, 1940). At present roe deer are evi- 
dently absent in this section although they did exist there even in the 1940's (B.V. 
Obraztsov). Thus roe deer appear and disappear periodically in this peripheral point 
of the geographic range. 

It is possible that the boundary of a fairly uniform distribution ran slightly 
more north and encounters at Novomoskovsk and the Black Forest of Kirovograd 
region are isolated cases. There are references (Korneev, 1952; and others) to the effect 
that the Siberian and not the European form of roe deer inhabited these two points. 
Verification is required however. 



269 

Bendera to Kaushan and farther to Oloneshta, dien turned southwest 
and going through Tarakliya ran across the Prut into Rumania. 
Thus the steppes of the Izmal' district do not form part of the 

162 

range. 

The outline of the western part of the Siberian-Middle Asiatic 
distribution of roe deer changed very extensively by the mid-1950's, 
moving considerably westward, especially in the southern part, 
quite significantly southward, and very extensively northward. The 
western boundary of this section of the range commenced in the 
south, on the Ural River slightly below the mouth of the Ilek (at 
Nizhne-Ozernaya), proceeded northwest to Totskoe, passed to the 
west of Buzuluk somewhere between Buzuluk and Kuibyshev, ran 
farther to Buguruslan, encompassed Belebeev rise from the west, evi- 
dently turned slightly east, and exited at Menzelinsk slightly 
southwest of the mouth of the Beloe. There the new boundary joined 
the reconstructed and later even coincided with it, or almost so. 
From Menzelinsk the boundary bent east or northeast fairly sharply 
and, in a manner not yet clearly understood, proceeded north along 
the foothills of the Urals somewhere between 57 and 58° ETong.'^'' 

The relatively small extension of the northern section of the 
western boundary into the west is explained by the very heavy snow 
in that section of the western Cis-Urals. There is essentially no 
possibility of further enlargement of the range into the west there, 
unlike in the south. The northward enlargement of the range of roe 
deer along the Urals has been very pronounced over the last decade. 
196 The outline of the range there corresponds to that described earlier 
in discussing the reconstructed range. There is, however, justifica- 
tion to assume that colonization of roe deer in the northern section 
of this region does not represent exclusively the restoration of the 
range which existed sometimes in the past, but also the occupation of 
new territory in which roe deer were previously absent. 

The southern boundary of this part of the range, commencing in 
the west in the Urals at the mouth of the Ilek, encompassed the Ilek 
basin together with Khobdo and proceeded farther through Kazakh- 
stan as described in the depiction of the reconstructed range. 

Visitations by roe deer to the mouth of the Kama are known from 
the area described here (V.A. Popov et al., 1954). 

'*^The boundary line described is based on data from Kalela, 1948; Fedosov, 1951; 
Danilov 1947; Siivonen, 1953; Bobrinskii, 1944; Korneev, 1952; Nasimovich, 1955; 
Formozov, 1946; Elpat'evskii, Larina and Golikova, 1950; Izmailov, 1937; Kuznetsov, 
1952; and others (including the unpublished data of B.V. Obraztsov, LP. 
Rasshivalina, and V.G. Heptner). 

^" According to Kuklin, 1951; V.A. Popov et al., 1954; and the unpublished data of 
V.A. Popov and S.V. Kirikov. 



270 

The advance of the eastern boundary eastward in the western 
area inhabited by roe deer, and the western boundary westward of 
the Siberian-Middle Asiatic region, indicate an eventual merger of 
these regions into a continuous range. The break in the 1920's 
extending for about 1,500 km or 24° had shrunk to 900 km or 10° 
longitude at the narrowest section (between 50 and 55° N lat.) by the 
mid-1950's. Roe deer visitations from west to east and vice versa, 
reaching the Volga, support this possibility. If appropriate conser- 
vation measures are undertaken, the merger of these parts of the 
range is entirely probable since the natural conditions are quite 

, • 164 

conducive. 

The contemporary southern boundary of the Siberian-Middle 
Asiatic distribution of roe deer coincides with the reconstructed 
range described before and represents, as pointed out, the boundary 
of the Middle Asiatic part of the range. The latter also corresponds 
to the range described above. 

The northern boundary of the range in Siberia and the Far East 
fully coincides with that of the reconstructed range all along its 
extent except for a small section beyond the Yenisey. There the 
boundary crosses the Yenisey slightly north of Krasnoyarsk, runs 
east roughly along 56°, intersects roughly at that latitude the upper 
courses of the Biryusa and Chuna,'>descends in the form of a promen- 
tory in the interfluve of the Chuna and Angara toward the south 
almost up to 55°, again rises in the north along the left bank of the 
Angara, and turning east crosses it at the mouth of the Ilim. From 
there it again turns south and, crossing the Lena at Zhigalov and the 
Kirenga almost at 55°, again rims north encompassing from the 
west the Northern Baikal and Patom highlands, crosses the Lena, 
and becomes the reconstructed boundary on the left bank of Lena in 
the Nyuya basin as described earlier. Thus, the present-day bound- 
ary lies somewhat south of the reconstructed one described 
above. '^^ 

As mentioned before, at some places in the region north of Bai- 
kal, especially in Yakutia, colonization of roe deer beyond the limits 
of the former range took place in the last decade. It should be 
emphasized that here the process proceeded extremely intensely, evi- 
dently more intensely than in other parts of the range. Judging from 
all available information the effect of natural causes was particu- 



'*" A merger of the Caucasian and, even less, the Crimean part of the range with 
the northern should not be anticipated. 

"' According to the data of Favorskii, 1936; Podarevskii, 1936; Kopylov, 1948 and 
1950; Fetisov, 1953; and other sources. Also see footnote no. 142. 



271 

larly important although the human factor (reduction of persecu- 
tion) was not negligible. All information relating to the last century, 
commencing from Maak (1859), points to the distribution of roe deer 
as a rare phenomenon in the north, only up to the mouth of the 
Vitim. One transgression was recorded on the Vilyuya toward Suntar 
in the region south of the southern bend of the Vilyuya. Slighdy more 
eastward deer were distributed along the Olekma. This was the 
situation until the last decade. According to informadon even for 
the 1920's, 1930's, and pardy 1940's the northern boundary of roe 
deer usually traversed south of the Northern Baikal highlands to the 
south of Bodaibo, and beyond in the lower reaches of the Chara and 
Olekma, and still farther into the passes in the Patom highlands 
from the south. Some data point to their occurrence along the Lena 
above the mouth of the Olekma. In other words, Yakutia along the 
left bank of Lena, and along the right bank east of the mouth of the 
Olekma was not inhabited by roe deer. 
197 An extremely intense colonization of roe deer took place in 
northern Yakutia in the 1940's and 1950's. During that period 
almost all the region between the Lena and the middle and southern 
Vilyuya was completely occupied by roe deer, which even crossed the 
northern Vilyuya. Colonization of roe deer in the region between the 
Lena and Amga, i.e., east of the Lena, commenced very recently, in 
1953 and 1954 (D.U. Ivanov).'^^ 

Colonization of roe deer in Yakutia continues and a further 
enlargement of the range there, especially north of the Vilyuya, can 
be expected. It is significant that roe deer occupied the area in Yaku- 
tia first along the Vilyuya and also along the Lena and the lower 
reaches of the Amga, all of which regions represent zones with natu- 
ral topographies of steppes and forest-steppe types. The thinning of 
forests in that part of the country was also favorable for these ani- 
mals. It is significant that at some places in the newly colonized locali- 
ties, for example around Yakutsk, roe deer prospered into fairly 
large populations (D.U. Ivanov). 

The present-day range of roe deer in the Crimea, as in the 1920's, 
is confined only to forest zones in the mountains of the peninsula. 

The range of roe deer in the Caucasus has not undergone marked 
variations since the time of maximum contraction, and whatever 



'" Some published maps of geographic ranges (Bobrinskii, 1944) in which the 
northern boundary along the right bank of the Lena runs from the Olekma in the 
form of an arc toward Uda Bay, terminating on the Aldan beyond 60° N long., do not 
at all represent the prevailing state in that period and were incorrect even for 1955 and 
1956. 



272 

changes occurred at some places were insignificant. This is explained 
by the occupation of roe deer, even in the 1920's, of almost all the 
area favorable for their survival, i.e., the forest zone of the Caucasus. 

The complex outlines of the range of deer in the Caucasus are 
connected with the forests. Commencing in the west around Novo- 
rossiisk, it extends in a belt along the forest zone of the northern 
foothills and the middle montane belt right up to Caspian Sea, 
running toward it in the area south from Makhachkala to Divicha 
(around 41° 10' N lat.). This band is extremely variable, depending 
on the width of the forest zone; in the west it is broad and reaches 
toward northern Maikop and Nal'chik; it is very narrow in the 
central part of the mountains and reaches only up to Vladikavkaz; it 
broadens again slightly in the east. Roe deer are absent in the whole 
of the high mountain region and in the forestless zones deep in the 
mountains, for example, in many regions of Dagestan. 

Along the southern slope of the mountains the range of roe deer 
extends in a very narrow band from the region of the source of the 
Pirsagat (north of Shemakha) up to Tbilisi and farther west. The 
range covers essentially the whole of the western Trans-Caucasus 
and extends in a narrow band along the southern slope of the Glav- 
noi range to the Sochi region where it joins the north Caucasian 
part of the range. All the steppe zone of eastern Trans-Caucasus up 
to Tbilisi inclusive is devoid of roe deer. They are also absent 
slightly west of Tbilisi along the Kur'. 

In the rest of the Trans-Caucasus the range is in the form of a 
narrow band running from the western Trans-Caucasus in the south, 
east toward southern Tbilisi, passing north of [Lake] Sevan and 
running farther southeast to Araksa. This band includes the moun- 
tain ranges of Papakar, Pambak, Shakhdag, Murov-Dag, and Kara- 
bakh. In the Zangezur mountains roe deer are evidently absent but 
their range includes the Kafan and Megra regions, associated with 
the Bargushat mountains, and the eastern slopes in the southern 
parts of Zangezur. Toward Araksa this projection of the range thus 
runs in the region between the Megra (southernmost point in the 
bend of the river) and the Karyagin region. 

In Armenia the southern boundary of this projection runs 
through Stepanovan and Kirovakan regions on the western margin 
of Sevan. North of Sevan this section of the range is very narrow and 
falls far short of the railroad and Kirovobad. In the region south of 
Tbilisi its eastern boundary extends approximately to Shamlug, run- 
ning west of the railroad to Shamlug. 

The bottomland deciduous forests of the lower Terek in the zone 



273 

198 slightly above Kizlyar and below the mouth of the Sunzha represent 
an altogether different section of habitation of roe deer in the 
Caucasus. Evidently roe deer are also encountered in the lower 
reaches of the Sulaka north of the railroad (Vereshchagin, 1947; map), 
where they were absent in the 1920's (V.G. Heptner). Another isolated 
section is the forest zone of Talysh, which represents an independent 
projection of the Iranian-southern Caspian part of the range of the 
species."^' 

In the last decade roe deer, mainly Siberian and Far East animals 
were transplanted at several places beyond the prevailing range: in 
Zavidov on the line connecting Moscow-Leningrad, on the southern 
bank of the Volga reservoir northwest of Klin (successful colonies, 
see above), on Pogonno-Losin Island near Moscow (this settlement 
was evidently unsuccessful), and in Prioka-Terras preserve near Ser- 
pukhov (successful). In the 1950's especially in the later half of this 
period, roe deer, mostly eastern forms (Siberian, Tien Shan, and Far 
East), have been colonized time and again at many places by game 
organizations. Sufficient information about these efforts is not 
available and in several cases the results are altogether unknown. 
These have therefore not been discussed here. 

Geographic Range outside the Soviet Union 

The range outside the Soviet Union (reconstructed) covered all of 
western Europe and in the south up to the Mediterranean Sea and 
Sicily inclusive (absent on other islands ). In the west the range 
covered England but not Ireland, and evidently never existed there. 
The northern boundary in the Scandinavian peninsula (including 
Finland) commenced on the Atlantic coast around 65° N lat., 
descended south, encircling the main axis of the Scandinavian moun- 
tains roughly up to 62°30' to 63°, and rose far north along the Pri- 
Baltic region of Sweden in the form of a band reaching the arctic 
circle (the northernmost point of habitation of the species). From 
there it descended steeply south and, traversing roughly along the 
eastern boundary of Finland, ran along the west coast of Lake 
Ladoga and the Karelian Isthmus.'"^ 

The range of roe deer in the Caucasus is based on data from Vereshchagin, 1947; 
Dal', 1954; Heptner and Formozov, 1941; and others (including original data of V.G. 
Heptner). 

The range boundary in Scandinavia is taken from Ekman, 1922; Kalela, 1948; 
and Siivonen, 1953. Roe deer regularly inhabited Sweden north only to approxi- 



274 

In the east the range covered Asia Minor, northern part of the 
Syrian zone of the United Arab Repubhc, northern Iraq and Pales- 
tine up to Lake Genisaret (Tiberius), and Haifa (Mount Carmel) 
and Iran. The range boundary in these countries to the south is not 
clearly understood. Evidently it extended in a narrow belt along the 
Elbruz and the adjoining forests of the southern Caspian district 
("Hyrcania"). In the east the range apparently penetrated up to the 
meridian of the southeastern corner of the Caspian Sea, but did not 
reach the Atrek basin. Roe deer were absent in the rest of Iran and in 
Afghanistan. 

In central and eastern Asia the range of roe deer included the 
eastern half of the Tien Shan, but the limits of their distribution in 
the east, in this part of China, are not clearly known. In Mongolia 
the range encompassed the Mongolian Altai roughly up to 92° E 
long., spurs of the Altai to the west of Lake Ubsu-Nur, Khan- 
Khukhei mountain to the northeast of Lake Khirgis-Nur, montane 
country of the Khangai in the south roughly to 46°, Khentei in the 
south to Ulan-Bator and almost 47° N lat., and approximately to 
115° E long, in the east (Bannikov, 1954). The Mongolian zone of 
habitation of the species led directly almost everywhere into the 
range within the USSR. 

Farther away in the east the range included a considerable part of 
the eastern half of China. The western boundary of this part of the 
199 range extended from west of the Great Khingan, including its spurs, 
to south of Lake Buir-Nur within the Mongolian Republic, and 
from the southern extremity of the Khingans, passing to the west of 
Peking, ran toward the Hwang Ho and the northeastern corner of its 
great northern bend. From there the range extended as a narrow 
projection into the west along the Inshan mountains lying north of 
the longitudinal course of the river. From the eastern extremity of 



mately 64° N lat. and the southeastern corner of Norway. The rest of the area repre- 
sented zones of fairly frequent or rare visits. In northern and northwestern Finland roe 
deer evidently originated from Sweden, and in southeastern and southern Finland 
from the USSR, mainly through Karelian Isthmus (migration through Svir dubious, 
V.H.). In Scandinavia in the present century intense colonization of roe deer in the 
north has occurred, especially in the last decade. The maximum contraction of the 
range took place at the end of the last century when roe deer occurred only in the 
extreme southern part of Sweden. In 1955 and 1956 they spread, as mentioned before, 
and not only reached their one-time maximum range, but at some places in the USSR 
even exceeded it. 



275 

the Inshan the boundary initially descended south and later ran 
west, bypassing the Ordos desert (within the arc of Hwang Ho) on 
the south. West of the Hwang Ho (that section running north) the 
range extended in the form of a narrow projection along the river to 
the north of the Ala Shan mountains. It also ran slightly west, 
occupying the eastern part of the Nan Shan range. Passing to the 
east of [Lake] Koko Nor the boundary turned south, including in the 
range eastern Tibet (regions of upper Hwang Ho, Yangtze, and 
Mekong) and the Kam country (Flerov, 1952). From there it became 
the southern boundary of the range, ran through northern Szech- 
wan, and covered southern Shensi and Shansi, approximately to the 
region of the Hwang Ho estuary. 

The range in China, namely in eastern and southeastern China, 
evidently changed very extensively long ago. It is highly probable 
that at some time in the past it extended to the Yangtze and occupied 
the entire counny between this river and the Hwang Ho. At present 
it is possible that roe deer inhabit the region in the northern bend 
region along the midcourse of the Yangtze. 

All of the Korean Peninsula and Cheju-Do Island fall within the 
range. 

Fossil remains of roe deer are known only from within the pres- 
ent-day range or at its boundaries (Syria — Pidoplichko, 1951; Kon- 
dopoga and right bank of the Svir — Kalela, 1948). (V.H.) 

Geographic Variation 

The magnitude of geographic variation in the species is significant 
from a purely morphological point of view, i.e., with reference to 
sharp differences between extreme forms. However, the number of 
races, considering the vastness of the range, is comparatively moder- 
ate and in any event much smaller than the number of names 
assigned to the species (see the synonyms). At the same time, in some 
limited areas, especially in western Europe, roe deer tend to show 
local features, mainly in size and weight, as a result of local ecologi- 
cal conditions, primarily food situation, population density, etc. 
There is no justification for considering these variations as factors 
dividing the species into subspecies. Some variations in this genus 
were even recorded here years back (differences between roe deer of 
western and eastern parts of the Great Caucasus; Dinnik, 1910). Such 
forms have generally not been recognized as subspecies and only 
prominent races are acknowledged in Soviet literature. At the same 
time geographic variability in species cannot be considered clearly 
understood since their range is obviously somewhat oversimplified 



276 

in the latest reviews (Ellerman and Morrison -Scott, 1951; Flerov, 
1952)."' 

In the USSR races of roe deer fall quite naturally into two dis- 
tinct groups — European and Siberian. 

European Group 

1. European roe deer, С с. capreolus Linnaeus, 1758 (syn. vulga- 
ris, caprea, europaeus, transsylvanicus, balticus, albicus, warthae, 
armenius, and zedlitzi). Small forms. Body length 100 to 136 cm, tail 
length 2.0 to 3.5 cm, height at shoulders 75.0 to 91.5 cm, maximum 
skull length 190 to 216 mm, and weight 20 to 37 kg'™ (Flerov, 1952; 
Sablina, 1956; Serzhanin, 1956). 
200 Winter coat grayish-brown, darker in rear part of trunk and 
much lighter on sides. Summer coat generally brownish-rusty or 
rusty. Top of head grayish or brownish, differing from color of 
trunk. Young animals rust-colored with bright spots arranged 
mainly in three rows. Spots in uppermost row sharper and brighter 
than the rest. Tympanic bulla very small, set deep in auditory 
region, projecting weakly above articular process (Flerov, 1952). 
Antlers poorly developed and characteristic of European type. 

Found in the European part of the USSR, including Belorussia, 
the Ukraine, Crimea, and Trans-Caucasus. Outside the USSR 
occurs throughout Europe, including the Balkans, Asia Minor, Iraq, 
Palestine, and Iran. 

Until quite recently, as partly revealed by synonyms, several sub- 
species were recognized in this territory. The description of the nom- 
inal form was published only recently (Ellerman and Morrison- 
Scott, 1951; Flerov, 1952). Although it reflects the prevailing state 
better than the former approach, it is nevertheless somewhat 
simplified. Thus it is doubtful if Trans-Caucasian and Iraqi roe deer 
are identical with southern Swedish, central European, and 
Belorussian animals. This subject requires special study. The name 
armenius Blackler, 1916 should perhaps be assigned to the trans- 
Caucasian form. 



Voluminous data on geographic variability of characteristics (mainly meristic) 
of roe deer have been given by Pasternak (1956); Unfortunately this information has 
not been thoroughly studied. Pasternak's findings differ somewhat from those given 
here. 

'™ The weight of 41 kg mentioned by Flerov (1952) evidently refers to a rare find. 
Latvian roe deer weigh on the average about 23 kg (Kalnin'sh, 1950) and German 
males 15 kg (Hempel et al., 1956). 



277 

Siberian Group 

2. Siberian roe deer, C. с pygargus Pallas, 1777. Largest form, 
attaining maximum size and found in the Urals, Pri-Baikal, and 
especially in the Altai-Sayan region. Body length of males 123 to 151 
cm (M 134) and females 119 to 147 cm (M 129), height of males at 
shoulders 84.5 to 100 cm (M 90) and of females. 80 to 96 cm (M 87.5), 
and height of males at sacrum 89 to 106 cm (M 98) and of females 87 
to 105 cm (M 96) (Pri-Baikal; Fetisov, 1953). Maximum skull length 
215 to 253 mm. Weight of males generally reaches 59 kg (Flerov, 
1952) and of females up to 52 kg (Fetisov, 1953). These animals are 
the largest of all the forms and possess the largest antlers in absolute 
as well as relative terms. 

Winter coat in general brownish or bluish. Speculum large and 
white. Summer color rusty or rusty-red without contrast between 
coloration of head, trunk, and neck. Spots well defined in young and 
arranged in four longitudinal rows with additional groups of spots 
occurring on shoulder or/and thigh. Spots in upper rows do not 
differ from lower ones in brightness. Skull in orbital region rela- 
tively slightly narrower than in European forms. Tympanic bulla 
large, swollen, and projects prominently in auditory region (Flerov, 
1952). Antlers in fully developed form, typical of Siberian type (see 
above). 

These animals occur in the eastern half of the European part of 
the USSR, the Urals, and Siberia up to the Amur. Outside the USSR 
they are known in northern Mongolia. 

It is very possible that roe deer are not taxonomically identical 
throughout the vast expanse from the Volga to Trans-Baikal. 

The range boundary of Siberian roe deer is quite vague in com- 
parison to the European since the roe deer is now extinct in all 
regions through which the [common] boundary once traversed. Evi- 
dently Siberian roe deer reached the Volga, occurring along its right 
bank, especially in the lower reaches of the Don (they lived in the town 
of Sarkel in the twelfth century; N.K. Vereshchagin). The boundary 
evidently did not run farther west of the Volga and much of the 
plains between the Volga and Dnieper were occupied by the Euro- 
pean form, which is established there even today. 

Reports that Siberian roe deer were encountered much farther 
201 west, at least in the historic past, are apparently erroneous and based 
on some individual animals with particularly large antlers found 
among European roe deer even in western Europe. 

Average body length of Altai-Sayan animals according to Pasternak (1955) 
140.4 cm; mean weight 35 kg. 



278 

Many reports exist that Siberian roe deer also lived on the right 
bank of the Dnieper in the Black Forest in Kirovograd region and on 
the left bank of the river in Samar forest in Novomoskovsk and 
Dnepropetrovsk regions (Brauner, 1915; Migulin, 1927 and 1929; 
Charlemagne, 1937; Korneev, 1952). Some regard this form there as a 
relict of the glacial period (Charlemagne, 1937), while others 
(Migulin, 1927 and 1929) concede transgressions into these places of 
Siberian or Caucasian animals. The subject requires special study, 
including a study of their systematics, but it is extremely doubtful if 
tiue Siberian roe deer lived in these places. This is particularly 
contraindicated by the occurrence of the European form in Crimea. 
Finds of remains of Siberian-type roe deer in Orlov region (Gorba- 
chev, 1915; Flerov, 1952) pertain evidently to extremely ancient 
periods. 

The foregoing discussion points up the near impossibility of 
delineating the range boundaries of both these forms between 30 and 
35°E long. (Flerov, 1952) over the historic period. It is entirely possi- 
ble that in the Quaternary period in the west, even in western 
Europe, roe deer of Siberian type did exist and that the western form 
represented a younger generation produced by diminution. How- 
ever, geographic relations over the historic period were probably the 
same as discussed above. 

At present it is nearly impossible to study the subject of former 
range boundaries since Siberian roe deer are not established west of 
the Volga, while the European type has noticeably advanced east- 
ward. Moreover, Siberian and Far East animals were introduced by 
man in the Moscow district and some other regions and the picture 
has been greatly confused. 

3. Tien Shan roe deer, C. с tianschanicus Satunin, 1906 (syn. 
jerganicus). Very similar to Siberian form in color but slightly 
smaller. Maximum skull length 218 to 238 mm (M 226). Features of 
similarity in the skull structure with the Caucasian form also evi- 
dent (Pasternak, 1955). Antlers large. 

Found in Tien Shan. Outside the USSR they occur in Chinese 
sector of Tien Shan. 

The type is not well known and the form within the Soviet 
Union has not been studied. Some authors place Tien Shan roe deer 
among Siberian forms but this is extremely dubious. 

4. Far East (Manchurian) roe deer, C. с bedfordi Thomas, 1908 
(syn. mantschuricus and ochracea). Very similar in structure to Sib- 
erian roe deer, from which it differs in slightly smaller overall size 
and smaller antlers. Body length averages about 122 cm and weight 
about 28 kg (Pasternak, 1955). Maximum skull length 211 to 236 



279 

mm and average 226 mm. Winter coat ocherous-rusty or gray with a 
rusty tinge. Speculum with faint rusty tinge. Summer coat bright 
rusty and full, more so than in Siberian form. 

Found in Ussuri region and Pri-Amur. Outside the USSR this 
form occurs in northeastern China and Korea. 

The range boundary of Far East roe deer in comparison with the 
Siberian form is not clear. Evidently roe deer of Great Khingan in 
the east, in expanses adjoining the Amur from the north and possibly 
in the Amur itself, possess transitional features from the Siberian 
form. In the most typical form this subspecies is found in China and 
the southern Ussuri region in USSR (Primor'e territory ).'^^ Thus, 
the northern range boundary of the subspecies and the region of its 
merger with the Siberian form fall in the USSR. 
202 5. North Caucasian roe deer, С с. caucasicus Dinnik, 1910. Size 
moderate, intermediate between European and Siberian roe deer. 
North Caucasian roe deer are fairly large and have antlers of the 
Siberian type. Body length reaches up to 1.5 m with an average of 
roughly 130 cm. Weight does not exceed 40 kg and heavy animals are 
very rare; average weight about 30 kg. Maximum skull length 216 to 
235 mm (M 225). Antlers not more than 33 cm in length. 

Found on the northern slopes of the Great Caucasian range from 
Novorossiisk and Sochi to the Caspian Sea coast and from the 
mouth of the Terek to Derbent. Does not occur outside the USSR. 

North Caucasian roe deer have been studied very little and their 
features are inadequately defined. Nevertheless, there is no justifica- 
tion for denying their independent existence and placing them 
among the true Siberian form (C. с pygargus; Flerov, 1952). In gen- 
eral the North Caucasian form is smaller than the Ural and other 
Siberian roe deer. 

Trans-Caucasian (C. c. capreolus) and North Caucasian forms 
are found together at some places (western extremity of the geogra- 
phic range, Dagestan, lower course of the Terek). Trans-Caucasian 
type animals, which cannot be distinguished from them, are also 
found within the range of North Caucasian forms. Apparently some 
show transgression of particular characters in both directions. 
There is no doubt of a fairly extensive hybridization of both forms. 
These relations have not been accurately delineated. It is possible 
that the race under description is generally of a hybrid character 
formed as a result of the encounter between two strains of Caucasian 

'Farther south, in the central and southern provinces of China and eastern Tibet, 
some authors acknowledge the existence of one more form of Siberian type close to 
the Far East type, i.e., C. с melanotis Miller. Others consider this form identical to C. с 
bedfordi.The latter view appeals more correct. 



280 

populations of roe deer, i.e., Siberian from the north and European 
from Asia Minor into Trans-Caucasus. In addition to these differ- 
ences, the North Caucasian form also differs distinctly from the Trans- 
Caucasian in skull measurements (skull length of roe deer from the 
latter area 184 to 213 mm— M 200) (Pasternak, 1955). 

The presence in northern Caucasus of Siberian-type roe deer 
serves as yet one more proof of the former occurrence of Siberian roe 
deer on the right bank of the Volga and along the lower Don. 

Outside the USSR, apart from the forms discussed above, yet 
another form occurs — C. с melanotis Mill., 1911 (northeast Tibet, 
Kansu, and Szechwan). It is proximate to the form bedjordi and 
possibly the two are identical. (V.H.) 

Biology 

Population. According to rough computations of data for the 1920's 
the average population density of roe deer in the forest areas of the 
USSR was 0.67 per 1,000 hectares (Lavrov, 1929). In the last few 
decades the range of roe deer has expanded and the animals have 
penetrated farther north into Siberian and Kazakhstan forest-steppes 
(see above) and multiplied. 

In the last decade roe deer were most numerous in eastern Sibe- 
ria, in the southern Far East, at some places in Kazakhstan forest- 
steppes, in the Altai, and Trans-Urals. In eastern Siberia roe deer 
were especially numerous in the southern Irkutsk and Chita regions 
and in Buryat and Tuvin Autonomous regions. Here their popula- 
tions rose in some places up to 25 to 50 animals per 1,000 hectares 
(Timofeev, 1949; Kopylov, 1950; Fetisov, 1953; and others). In the 
Far East roe deer were most numerous in the Jewish autonomous 
and Amur districts (Adramov, 1954). In the Altai they were particu- 
larly numerous in the Gorno-Altai Autonomous Region, especially 
in Ust-Kan, Ust-Koksin, Shebalino, and some other regions. In 
Il'men preserve, from the end of the 1930's to the early 1940's, over 
an area of about 50,000 hectares there were usually 1,200 to 1,500 roe 
deer in winter, which in the exceptionally snowy winter of 1940 to 
1941 rose to 3,500 (Ushkov, 1954). 

In the European part of the USSR roe deer are a common sight in 
the Baltics (mostly in Lithuania and Latvia), Belovezh Forest, Cri- 
mea, and at places in the Caucasus. In Latvia up to 34,000 roe deer 
were counted in 1939 but their number decreased in the years follow- 
203 ing the Great Fatherland War and the population was restored later: 
8,500 animals in 1948; 15,500 in 1955; and 17,880 in 1957 (Ivanaus- 
kas, 1957). 



281 

In Crimean preserve 1,300 roe deer were counted in 1950, or an 
average of about 40 animals per 1,000 hectares in the area in which 
the animals lived. Outside the preserve in Crimea the population 
touched 3,300 animals, with the density being highest in the Alush- 
tin region (35 per 1,000 hectares), Staro-Crimea (28), Bakchisarai 
(20), etc. (P. A. Yanushko). In the early 1920's there were about 5,000 
roe deer in Belovezh Forest (Severtsov, 1940; and others). Later, the 
population there decreased to about 500 by the end of the 1940's (by 
that time the area of the Forest was also reduced); in some individual 
woods density varied from 1 to 18 animals per 1,000 hectares (Sab- 
lina). In the Ukraine and Belorussia on the whole, as also in the 
central regions of the European part of the USSR, roe deer were few.'^^ 

In the mid- 1 800 's the population of roe deer was markedly high 
almost everywhere. It was exceptionally so in the Urals, southern 
Siberia, and the Far East, where herds of several hundred animals 
were sighted during seasonal migrations (Sabaneev, 1875; Nasimo- 
vich, 1955). 

Habitat. Roe deer inhabit regions with diverse climates, from 
humid marine to dry continental. They inhabit forest-steppe sec- 
tions and diverse types of deciduous, mixed, and coniferous forests, 
as well as tugais, thickets of reeds or rushes along the periphery of 
steppes, and sometimes areas around semidesert lakes. When 
rational game management is practiced, they survive well in culti- 
vated places and densely populated areas. In the snowfree period of 
the year the vertical range of dispersal is quite extensive, from sea 
coasts to subalpine and lower alpine meadows; in the Urals and 
Trans-Baikal up to 1,000 to 1,500 m (mountain-steppe meadows and 
balds); in the Caucasus and Altai up to 2,000 m; and in the Tien 
Shan mountains up to 2,800 to 3,300 m. They prefer smooth areas in 
the mountains and are more common in rugged localities in the 
plains and foothills. They prefer light forests with abundant young 
deciduous trees and undergrowth alternating with glades, clearings, 
and meadows; they avoid dense boreal forests of the taiga type. The 
extensive penetration of roe deer into the taiga zone in the last 
decade was facilitated by the felling of dense coniferous bottomland 
forests and the creation of burned-over forests with reseeded decidu- 
ous tree species (Skalon, 1946; and others). In the past roe deer of the 
forest-steppes penetrated far into the open steppes and even today 



'"in cultivated lands, especially when the population of competing species is low, 
the population density of roe deer can run very high; for example, in Ba\aria it 
averages 84 per 1,000 hectares (Grushvits, (1957).* 

*Not in Literature Cited — Sci. Ed. 



282 

are seen at some places in western Siberia and Trans-Baikal under 
similar conditions, but in very small numbers. 

The preference of roe deer for a given type of habitat varies 
widely in different sections of the geographic range. For example, in 
Belovezh Forest roe deer are particularly numerous in mixed pine- 
oak-spruce forests, among pine undergrowth, and in oak-hornbeam 
and beech forests; roe deer also visit marshes (Sablina, 1955; Ser- 
zhanin, 1955). In Lithuania roe deer inhabit forests of all types and 
composition but are mostly confined to mixed forests with well- 
developed young trees, burned-over forests, and clearings; they are 
scarce in spruce and even scarcer in pine groves (Kuznetsov, 1954). In 
the Caucasus, although roe deer are encountered in summer right up 
to subalpine meadows, they are most numerous in broad-leaved 
forest sections with young deciduous varieties of trees and large 
clearings covered with dense herbaceous vegetation seeded by man 
(Dinnik, 1910; Nasimovich, 1939). Beech forests with tall trunks and 
numerous young trees represent one of the favorite summer habitats 
of roe deer in Crimea (Lavrov, 1929; Dal', 1930). In the Urals and 
western Siberia roe deer prefer mixed pine-birch forests with aspen 
groves, 10-year-old burned-over forest sections and clearings, and are 
204 attracted to plantations adjoining glades and marshes, common in 
the floodplains of rivers. In the steppes they are confined to forest 
islets and reed thickets along the banks of lakes (Sabaneev, 1875; 
Vinitskii, 1937; Averin, 1949; Kurazhskovskii, 1949; Yanushevich 
and Blagoveshchenskii, 1952; LP. Laptev, 1958). 

In the Altai, eastern Siberia, and along the Amur roe deer are 
particularly numerous in deciduous (at places also pine) forests with 
dense deciduous undergrowth of shrubs and grasses, in regenerating 
burned-over forests, in tiny birch-aspen groves adjoining cultivated 
fields, meadows, and steppes (Altai, the Sayans, and Trans-Baikal). 
Roe deer avidly feed in sedge-clover marshes,* sorghum fields 
(Altai), open ridges, and burned-over forests. They seek refuge in 
boreal forests, shrubbery thickets along river banks, etc. (Cherkasov, 
1884; Dmitriev, 1938; Nasimovich, 1949; Pavlov, 1949; Timofeev, 
1949; Kopylov, 1950; Fetisov, 1953). In the Far East roe deer are 
confined to sparse deciduous and burned-over forests; they penetrate 
deep into the Sikhote-Alin only along river valleys and avoid coni- 
ferous forests (Kaplanov, 1948; and others). 

The habitats of roe deer are extremely diverse in Middle Asia, 
where this species is common in spruce forests (central, northern, 
and eastern Tien Shan), often encountered in fruit- and nut-bearing 

*Not clear; may mean three-leaf sedge marsh — Sci. Ed. 



283 

forests in midmontane altitudes, juniper forests, along tugais in 
rusfies close to water sources, etc. In summer roe deer are most 
numerous in the subalpine belt of the mountains of central and 
eastern Tien Shan, where areas of spruce alternate with shrub 
thickets and tall grass meadows; the animals ascend to alpine mea- 
dows during early morning hours (Shnitnikov, 1936; Vakulenko- 
Snigirevskaya, 1940; Antipin, 1941; Kuznetsov, 1948; Shul'pin, 1948; 
Pasternak, 1955; and others). In the Tersk Alatau roe deer are found 
even in winter at heights of up to 3,200 m (R.P. Zimina). 

Food. In the Soviet Union the food of roe deer has been best 
studied in Il'men preserve (Trans-Urals) (Averin, 1949), Irkutsk 
region, Trans-Baikal (Fetisov, 1947 and 1953), Belovezh Forest (Sab- 
lina, 1955; Pasternak, 1955), Crimea (Lavrov, 1929; Dal', 1930; P.A. 
Yanushko), and the Baltics. Observations are too inadequate in 
other regions to draw justifiable conclusions. Two relatively impor- 
tant food components for roe deer are herbaceous and tree and shrub 
vegetation; their relative importance varies according to topographic- 
geographic characteristics of a zone and its natural features. For 
example, in the Crimea, throughout the year including even 
summer, tree and shrub vegetation is of utmost importance in the 
diet of roe deer, while herbaceous and bulbous plants are important 
in other steppes and semideserts, and twigs in winter. 

Usually leaves and green shoots of tree and shrub vegetation are 
consumed in summer and in winter shoots, branches, buds, and dry 
leaves (especially of aspen), and coniferous needles (most often of 
young pine, later juniper, and very rarely larch, spruce, or fir; nee- 
dles of nut pines are not eaten). Tree bark is not usually touched and 
roe deer differ significantly in this respect from red deer and elk. In 
some regions (Belovezh Forest and Bashkir preserve) the importance 
of twigs in the diet of roe deer is more than in the diet of red deer 
(P.F. Kaznevskii). Among herbaceous vegetation roe deer consume 
avidly tips of shoots, racemes, and sometimes syncarps [infructescen- 
ces] (western Siberia; Kurazhskovskii, 1952); in winter green parts 
and twigs projecting above the snow cover are also eaten. Black- 
berry, mistletoe (in forest clearances), winter horsetail, ivy, moun- 
tain oats (in northern Caucasus), and some other food plants are 
very important among winter-green plants. 

Qn the whole roe deer of the Soviet Union consume up to 250 

species of plants. In Crimean preserve the consumption of 145 

205 species of plants has been established (100 species of herbaceous 

According to Danilov (1958) roe deer consume about 174 genera of angiosperms 
alone (41 tree and shrub genera and 133 herbs). 



284 

plants, 29 tree and shrub, 4 lichens, etc.); of these, over 100 species 
are consumed in summer (including 63 which are eaten more often) 
and in winter 41 (and 14) (P. A. Yanushko). In the Irkutsk region and 
Trans-Baikal roe deer consume no less than 130 species (including 
85 eaten more often) (Fetisov, 1953). In Il'men preserve 110 plant 
species are consumed by roe deer; of these, in spring 90, in summer 
83, and in autumn and winter 64 or 65. Of primary or secondary 
importance are 24 plant species in spring, 34 in summer, 28 in 
autumn, and 15 in winter (Averin, 1949). In Belovezh Forest roe deer 
consume over 90 species (of which roughly 70% are herbs). In 
summer, compared with winter, food composition is far more 
diversified (Pasternak, 1955; Sablina, 1955). Plants serving as food 
for roe deer in the Altai include about 60 species (Pasternak, 1955). 

Among tree and shrub foods preferred by roe deer and which 
usually play an important role in their diet are aspen, ivy, birch, 
linden, mountain ash, oak, ash, some maples, hornbeam (in the 
European part of the USSR), hazel nut, raspberry, etc. Several spe- 
cies, mainly because of their extremely limited availability, are 
important only in small parts of the geographic range, for example, 
dogwood and jasmine in the Crimea and the Caucasus, smoke tree 
in the Crimea, Daurian rhododendron in Trans-Baikal, and lesp- 
edeza, Amur vine, actinidia, etc. in Primor'e. At places where the 
population of roe deer or tree-eating ungulates is high and preferred 
foods inadequate, roe deer consume large amounts of pine needdles in 
winter (Il'men preserve, Belovezh Forest, Baltics, etc.). Pine needles 
are usually most intensely consumed in the latter half of winter 
when the height of the snow cover is considerable (Averin, 1949). 
Thus pine food may serve as a fairly reliable index of winter food 
availability for this species. The same can be said of spruce with 
even greater justification; spruce is usually consumed only when the 
population density is extremely high, as observed in the Baltics. 

Among herbaceous plants of great importance in spring are 
umbellifers (cow parsnip, angelica, and chervil), composites (yar- 
row, hawkweed, and crepis), legumes (pea vine, vetch, and clover), 
buttercups (double buttercup, crowfoot, and anemone) — and in early 
summer, sedges and grasses (new growth is consumed avidly, as is 
old). Avidly consumed species also include some gentians, especially 
buckbean, in search of which roe deer often visit marshes, geranium, 
bistort, St. John's wort, burnet, and in southern Siberia lilies (bulbs 

" However, in the Naryn region of Alma-Ata district where roe deer are few, they 
quite frequently consume spruce needles in winter; in one case these needles con- 
stituted up to 50% of the stomach contents (Zhirnov, 1958). 



285 

of Allium, martagon, and yellow lily), etc.; in spring pasque flower 
(Pulsatilla patens), Scilla, lungwort, primula, and others are also 
eaten well. Many plant species consumed by roe deer in large quanti- 
ties are deleterious to cattle. 

Roe deer avidly consume fungi (armillaria, pepper mushroom, 
brown mushroom, chanterelle, and others), but these are not signifi- 
cant in their diet. In winter arboreal lichens (five to seven species) 
are often consumed avidly. In many regions acorns, fruits of beech 
and other fruit-bearing trees (wild pear, apple, and others) serve as 
important food; birch catkins and many berries (mountain cran- 
berry, blueberry, bilberry, cloudberry, bearberry, etc.) are consumed 
avidly. Fruits of nut pines, chestnut, sea buckthorn, and "birch lips" 
are also eaten (Cherkasov, 1884). 

In winter, when necessary, roe deer can dig up twigs, etc. and 
acorns from under snow 10 to 15 cm deep. Instances are known of 
their digging up aspen leaves from under snow 1.5 m deep. Instances 
are also known of roe deer digging up and consuming fallen larch 
206 needles (Trans-Baikal). In their diet selection roe deer are more adap- 
table than red deer in some regions (Yanushevich, 1950; Sablina, 
1955), and inferior to the latter in other zones. 

In most regions roe deer move well to natural and artificial salt 
licks. In Primor'e territory they emerge on the sea coast in search of 
aquatic algae and other marine products covered with salt 
(Abramov, 1954); in winter they eagerly visit ice crusts, including 
those in marshes, and lick them (Cherkasov, 1884; Fetisov, 1953). 

Home range. Throughout the year, except for autumn and 
spring migrations, herds and solitary roe deer are confined more or 
less to definite home ranges, but when their populations increase to 
a considerable size their home ranges overlap one another. Observa- 
tions have been made in Slovakia of the active defense of territory by 
some males outside the period of rut against other males (Dyk, 1956). 
Home range size is greater in transitional seasons (spring and end of 
summer to early winter); in autumn even resident roe deer wander 10 
to 20 km a single day within a home lange of up to 10 km in 
diameter (Sabaneev, 1875; Ushkov, 1947). With snowfalls and an 
increase in snow level, the radius of daily travel decreases, becoming 
least in the latter half of winter and in the period of snow crusts. At 
this time of year some herds of roe deer live for a few days or even a 



"* Instances are known of captured roe deer with their stomachs filled predomi- 
nantly with pasque flowers, which are deleterious to cattle (Fetisov, 1953). 

In the Crimea acorn shells sometimes form up to 30% of the weight of the 
stomach contents of roe deer in winter (P. A. Yanushko). 



286 

week in an area of 0.3 to 2.0 hectares (western Caucasus, Belovezh 
Forest, and Trans-Urals).^^* In summer, except for the period of 
fawning and the first one or two weeks after it, roe deer wander 
much more widely than in winter. When water supply is inade- 
quate, they walk several kilometers in search of a water source 
(Naurzum pine forest). 

Daily activity and behavior. In summer roe deer graze in the 
morning, evening, and at night, but at intervals (Averin, 1949; 
Fetisov, 1953). When the day is hot, they rest and ruminate, often 
lying in dense grass, among ferns, bushes, or wind-felled trees, eat 
grass or leaves only in the proximity of the bed and usually in small 
amounts. In cloudy and cold weather they often feed during the 
daylight hours. They seek refuge from blood-sucking insects in 
rocky open forests, on windblown banks of lakes, and in dense 
vegetation. 

In winter roe deer graze at any time of day but if frequently 
threatened by hunters lie low during the day, emerging stealthily 
when necessary into very dense sections of forests, glens, etc., and 
coming out at night to feed in swampy river valleys (Altai), etc. 
During blizzards they seclude themselves in forest thickets. 

Roe deer can jump to a height of 2.0 to 2.5 m. In the warm period 
they sometimes bathe and swim well; during migration they freely 
swim through such rivers as the Angara, Yenisey, Ussuri. and Amur. 
The weight distribution on the hoof is high, 300 to 360 g per cm , as 
a result of which roe deer get stuck in deep snow. The critical height 
of snow cover for European, Caucasian, and Middle Asiatic roe deer 
is 30 to 40 cm and for Siberian roe deer 40 to 50 cm. On experiencing 
difficulties in movement on snow roe deer often use their own old 
tracks or tracks made by other ungulates to reach main thoroughfares. 

Smell and hearing are well developed in roe deer; in detecting 
danger they are guided to a lesser extent by vision than by olfaction 
and audition. 

In summer adult roe deer are solitary (females move with their 
young) and only rarely are two or three (usually males) encountered 
together (up to seven in exceptional cases). At the end of rut, in 
September-October, mixed groups (males, females, and young) 
sometimes form, consisting of 20 to 30 animals or more. At the 

"^ According to Fetisov (1953) and Dul'keit (1956) solitary animals and herds of 
roe deer live within a home range of 1.0 to 6.0 sq km throughout winter. 

"' In the Urals in the mid-1800's herds of roe deer comprised as many as 50 to 100 
or even 500 animals in the period of autumn migration (Sabaneev, 1875); in the 
snowy winter of 1940 to 1941 groups of roe deer numbering 50 to 120 animals were 
sighted in Il'men preserve (Ushkov, 1954). 



287 

conclusion of autumn migrations these groups usually break up 
into smaller ones consisting of two to four each or slightly more. In 
spring, in April, with the approach of the fawning season, even 
these groups usually split up. Thus the gregarious tendency of roe 
deer is maximum in autumn, followed by winter; to some extent, it 
is directly proportional to the total population of roe deer and the 
degree of snowiness of a given winter. The herd composition does 
not remain constant even for one month, let alone two; groups of 
over 10 to 15 animals split up rather readily. 

Seasonal migrations and transgressions. Seasonal migrations are 
characteristic of roe deer in many parts of the range. The course of 
these migrations is determined largely by the snow cover regime and 
the food regime. Migrations occur in autumn into less snowy 
regions and in spring into summer pastures and are particularly 
characteristic of forest-steppe zones (southern Trans-Urals, Kulun- 
dinsk, Minusinsk, and Achinsk steppes, Trans-Baikal, etc.), the 
Amur basin (in the past even m the Ussuri), and some northern parts 
of the range, but only in those areas where roe deer are fairly numer- 
ous (basins of the Chulym and Yenisey Rivers, Pri-Baikal, and 
Vitim plateau). Extensive migrations are common in many montane 
regions with relatively high (over 40 to 50 cm) snow cover (the Cau- 
casus, Altai, Sayans, and others). The spatial magnitude of the sea- 
sonal migrations of roe deer goes up (at least it did in the recent past) 
in the Amur basin to 200 to 300 km [sic], and in the Urals, Abakansk 
range, Kuznetsk Alatau, and western Trans-Baikal to 100 km or 
more. A few centuries ago seasonal migrations of roe deer into places 
which combined different topographic sections and zones were char- 
acterized by an even broader spatial magnitude and were observed in 
a much larger number of regions in the country, for example in the 
Ukraine (Nasimovich, 1955). In forests of the European part of the 
USSR and at places in other parts of the range seasonal migrations 
of roe deer are presently poor and involve displacement from one 
type of habitat to another within a few kilometers. 

Forest sections with abundant young deciduous trees and under- 
growth, low-snow steppes (especially in Trans-Baikal and in Minu- 
sinsk depression), '^° lower less snowy mountain altitudes, and 
generally places in which winter conditions are not very severe serve 
as places of winter congregation of roe deer. The range of vertical 
migrations usually does not exceed 500 to 800 m. In some montane 



'*" Roe deer are confined only in the warm period of the year to steppe zones where 
the snow regime is more severe (Kulundinsk steppe), and move away into forest sec- 
tions in winter. 



288 . 

regions roe deer spend part of the winter even at a higher level than 
in summer by congregating along windblown areas (Trans-Ili Ala- 
tau and Kungei Alatau). 

Migrations of roe deer to wintering sites occur in different parts 
of the range from mid-September to January and continue for three 
weeks to two months or more. Autumn-winter migrations usually 
commence with the first snowfall, particularly if the snow cover 
forms immediately, and rarely with the onset of sharp cooling. In 
mid- September, migrations commence in the Kuzetsk Alatau moun- 
tains, in eastern Sayan, Vitim plateau, and Zeya basin; the last 
migrations take place in December in the mountains of the Trans-Ili 
Alatau. In most of the remaining regions in the Soviet Union migra- 
tions of roe deer usually commence in October and end in November, 
usually earlier than in the case of elk and Manchurian wapiti. Dur- 




207 Fig. 64. Regions of winter congregation of roe deer from the 1860's to the 
1870's in the Trans-Urals in the Iset River basin and its uibutaries (scale in 

km). 

l_course of travel of roe deer to wintering sites; 2— region under influence of snow; 

3— places of winter congregation of roe deer. Scheme based on data of Sabaneev, 1872 

(from Nasimovich, 1955). 



289 

ing migrations roe deer move in groups, sometimes in tens; herds of 
100 to 500 animals have been recorded in the past. Spring migrations 
occur in April-May within a much reduced period (Nasimovich, 
1955). A few decades earlier an immense number of roe deer moved 
away in autumn and early winter from the left bank of the Amur 
into Manchuria; in most places roe deer cross the Amur only on ice. 
The points of crossing of migrating roe deer over rivers (on ice and 
by swimming) are usually fixed. 

The distance covered and vertical range of seasonal migradons 
also depend on the snowiness of a given winter. For example, in 
Crimean preserve, judging from the relative frequency of roe and red 
deer, only 5 to 7% of the summer population of roe deer in less snowy 
winters move out of the sanctuary down the slopes, while over 60% 
do so in highly snowy winters (P.A. Yanushko). At some places, in 
the latter half of winter, after a perceptible rise in the height of snow 
cover, fresh movements of roe deer are observed, which are some- 
times more significant than those at the commencement of winter 
(the Caucasus, southern Trans-Urals, left bank of the Amur, etc.). 

In Middle Asia and sometimes in the Caucasus individual roe 
deer emerge in summer into high mountain meadows, and in south- 
ern Siberia balds. Instances are known of migrations of roe deer 
during summer droughts; for example, in the summer of 1951 many 
roe deer from Belovezh Forest preserve, hit by drought, migrated 
into the territory of adjacent Poland (S.S. Donaurov). 

Reproduction. In most regions mass rut in roe deer occurs in the 
latter half of summer, sometimes covering early autumn (up to mid- 
September), but usually lasting no more than a month in a given 
year. The earliest period of commencement of rut is mid-July, but 
even earlier in certain years; this is especially true of western parts of 
the geographic range (western Europe, '^^ Baltics, western Ukraine, 
and Belorussia) and evidently also of Primor'e territory. In Latvia 
rut in roe deer sometimes commences from the end of June; in rare 
instances it has even begun from the middle of that month (Kal- 
nin'sh, 1950). Mass rut in roe deer commences from the end of July 
to early August in the Crimea, Caucasus, southern Urals, in Trans- 
Urals and most regions of Middle Asia, i.e., in regions with 
extremely diverse climatic conditions. The very last period for the 
commencement of mass rut is August 5 to 15 and is typical of west- 
209 ern and eastern Siberia (plains of western Siberia, the Altai, Sayans, 
Pri-Baikal, Trans-Baikal, Hentei in Mongolia, Yakutia, and else- 
Excluding montane regions where rut usually commences two or three weeks 
later. 



290 

where), i.e., regions with a definite continental cHmate. There is 
almost no doubt that this relates to the duration of winter and later 
to a cold spring, which is unfavorable to early fawning. Thus, from 
west to east (excluding Primor'e) the period of commencement of 
mass rut is, in general, steadily delayed. 

Rut usually terminates in the latter half of August and early 
September in western sections of the range (including Baltics, the 
Ukraine, and Belorussia), Crimea, Primor'e, and at places in the 
Caucasus and Middle Asia. In the Urals, western and eastern Sibe- 
ria, and at places in the Caucasus (Armenia) and in Middle Asia 
(Kungei Alatau) the period of rut extends up to mid-September, and 
in some years even up to the last ten days of that month or even early 
October (Pri-Baikal, Trans-Baikal, and Yakutia). In Crimean pre- 
serve mating was observed once in December (Dal', 1930); such 
instances in October-November are known for many regions. ^^^ 

Available data are inadequate to draw any distinct geographic 
patterns of the extent of regularity in the period of rut of roe deer in 
different regions. The commencement of rut, its intensity, and over- 
all duration in different years depends on a combination of several 
factors: the nature of the preceding winter and spring (in arid 
regions also the type of summer in the preceding year), weather 
conditions in the summer of a given year, and other conditions 
which determine the general well-being of the animals. At places 
where these conditions are most constant year after year the com- 
mencement and duration of rut are also most constant. 

Roe deer become sexually mature in their second year, but many 
breed only a year later; this is particularly true of males which other- 
wise cannot withstand successfully the competition from older and 
more powerful rivals. Young animals breed later than older ones. 
Among adult females as many as 10 to 20% may remain barren. 

During the period of rut males are extremely high-strung, cast 
off their usual caution (instances are known of their attacking 
men ), and emit a chuckling sound. Combats between males are 
frequent and sometimes end with injury to, or even the death of, one 
or both competitors. In the period of rut a male lives for a few days 
with a single female, mating with her several times. In rare instances 
two or even three males may cover a single female during the same 
period (Sabaneev, 1875; and others) or a single male cover two or 
three adult females. The male chases away the offspring, which 

'References by Sabaneev (1875) to instances of mating in March between young 
roe deer in the Urals beg credulity. 

Such instances also occur at other times of the year. 



291 

rejoin the female only after rut terminates. The female remains in a 
state of estrus for four or five days (Kalnin'sh, 1950). As soon as a 
female becomes unresponsive the male runs after a second and some- 
times even a third. Roe deer therefore exhibit facultative polygamy. 

The pursuit of a female by a male usually occurs in a clearing or 
the edge of a forest. The female endeavoring to escape her unwanted 
suitor often circles round a tree, boulder, stump, etc., as a result of 
which the turf and grass there are trampled down. Instances are 
known of rutting males inflicting mortal injuries on females and 
young deer (Averin, 1949; and others). Males in rut eat little and lose 
considerable weight. 

The fertilized egg travels along the oviduct for a few days, under- 
goes cleavage, and enters the uterus. There it remains in a state of 
relative inactivity, cleavage progresses slowly, and blastocyst forma- 
tion is delayed roughly up to December. Only from the end of 
December or in mid-January is the fetus perceptible to the naked 
eye. The commencement of active growth of the fetus is sometimes 
210 accompanied by excitability, similar to the behavior of martens dur- 
ing pseudoestrus. Among females mating in November-December 
delayed implantation is usually absent and embryonic growth com- 
mences immediately; fawning usually occurs at the same time as in 
females which mated in the latter half of summer (Lavrov, 1929; 
Kalnin'sh, 1950; Kirshenblat, 1951; and others'^"). Thus gestation 
can vary from 6 to 10 months; in most cases it extends over slightly 
more than 9 months (about 40 weeks). 

In the western part of the range, as in the Crimea and Caucasus, 
mass fawning occurs in May. In some years some females fawn at the 
end of April, while others do so only in the first half of June. In 
the Urals, western and eastern Siberia, and also in most Middle 
Asian regions mass fawning occurs two or three weeks later, com- 
mencing in the latter half of May to early June and ending around 
June 15 to 20; compared with the western parts of the range (Crimea 
and the Caucasus), fawning here occurs more uniformly. The young 
appear in Primor'e territory roughly around the same period as in 
Siberia. 

In some instances roe deer give birth much earlier or later than the 
period indicated above. In Crimean preserve instances of fawning 
have been registered on April 7 (1950) or even on March 1 (1948) 



The main patterns of egg development in roe deer were first studied by Bischoff 
(1854). 

In the western parts of the Ukraine some females give birth to young even by 
early April (Tatarinov, 1956). 



292 

(P. A. Yanushko), in the Irkutsk region in the first ten days of July 
(Fetisov, 1953), and in Il'men preserve on July 14, 1938'^^ (Averin, 
1949). 

In rare instances up to five (Sablina, 1955) or even six (Cher- 
kasov, 1884) embryos have been detected. Some were evidently 
resorbed or the young stillborn since instances of even four fawns 
wath a single female are exceptionally rare (observed in the Crimea, 
Il'men preserve, and elsewhere). Roe deer in most regions of western 
and eastern Siberia often bear two fawns, rarely one, and even more 
rarely three. Instances of triplets there are notably more frequent 
than in other parts of the range where climatic conditions are less 
severe. One or two fawns are usual in the Urals, Middle Asia,'^^ 
the Caucasus, Crimea, and at some places in the western parts of the 
range where the fertility of roe deer is usually somewhat less. Thus, 
of the 92 female roe deer with fawns counted in Crimean preserve in 
the summers of 1947 to 1950, 60% had one fawn each and 40% two 
each (deer with three or four fawns were not seen in those years) 
(P.A. Yanushko). In Belovezh Forest 169 roe deer were marked with 
young in the summers of 1945 to 1949; of these, 57.5% had one fawn 
each, 41.6% two each, and 0.9% (one deer) three (Sablina, 1955). In 
Il'men preserve 36 roe deer were found with fawns from May to July, 
1938 and 1939; 50.9% had one fawn each, 43.3% two each, 3.9% (two 
deer) three, and 1.9% (one deer) four (Averin, 1949). Thus the average 
number of fawns per mother deer in these parts of the range for the 
summer period was 1.3 to 1.4. 

Growth, development, and ynolt. Newborns remain stationary 
for a week, concealed in the grass. In the case of two or three fawns 
they do not remain together but stay at some distance from each 
other. At the age of six to eight days they begin to move away from 
the mother. At the end of two weeks fawns are not inferior to their 
mother in their ability to run. 

Lactation continues for two or three months and ceases just 
before the onset of estrus. The latest date for lactating females is 
211 October 20 (Fetisov, 1953). The young commence grazing on grass at 
the age of one month. In the period of estrus fawns remain aloof and 
join their mother later, after which they move together until the 
following spring. 

'^* The first fawns in that year were observed on May 10. Thus the overall duration 
of fawning exceeded two months. 

'^' According to Sabaneev (1875) three embryos used to be common in the Urals, 
while instances of females bearing four were not so rare. 

'** Instances of triplets have been recorded in Narynkol' region of Alma-Ata dis- 
trict (Zhirnov, 1958). 



293 

Two fully formed fetuses (males) covered with hair were removed 
from females caught in Belovezh Forest on July 7; they weighed 950 
and 1 ,230 g. In the first half of July fawns reach a weight of 5.5 kg or 
slightly more (Sablina, 1955). Newborn roe deer are exceptionally 
light in coloration and their spots not as sharply defined as at the 
age of two or three months when a coarser beard [guard hairs] forms 
and the background color becomes darker. The spotted juvenile coat 
molts into the unspotted adult coat during August or at places even 
from the end of July. 

Roe deer are born with all their milk incisors and premolars. 
Loss and replacement of milk incisors with permanent teeth take 
place at the age of 5 to 12 months; milk premolars are replaced with 
permanent ones at the age of 13 or 14 months. Molars begin to 
appear at the age of three months. In winter the rate of dental 
growth slows down but intensifies again from May; teeth are fully 
formed by the age of 15 months (Sokolov, 1956). 

Antlers become visible in young males from the autumn of the 
first year in the form of small "pipes"; growth ceases only by April 
of the following year. Usually (but not always) they have a single 
beam without tines. The first antlers are shed in December and new 
ones become evident in January, which keep growing until spring; 
by then, the total number of tines on each antler is two or three. 
Often, even by the third year, antlers attain full development with, 
usually, three tines each. According to observations made in the 
Baltic region roe deer antlers grow best in areas where the soil is rich 
in phosphates and calcium salts (Kalnin'sh, 1950). Among adult 
males antlers grow from the end of March to early May. Shedding of 
velvet ceases by May-June, ossification is completed a little later, 
and antlers are shed in November (at places from the end of October 
and just before December^*^). In young males, compared with older 
ones, antlers grow roughly two to four weeks later and sometimes 
only in June in the Crimea (Dal', 1930). Antlerless males and 
females with antlers are not uncommon. Deformed antlers are also 
not infrequent. 

Roe deer molt once a year, in spring. In January-February the 
hair is perceptibly worn out and shedding commences in March- 
April, initially from the sides of the trunk below the shoulder blades, 
later from the legs, head, and neck, and last of all from the rear of the 
back. Depending on the condition of spring, molt usually peaks the 

'*' Instances are known when roe deer shed their antlers even in September (Cri- 
mea; P.A. Yanushko) and, contraiily, when antlers were still present even in early 
January (Irkutsk region; Fetisov, 1953). 



294 

latter half of April to early May and ends in June and in rare cases in 
early July. In the montane regions of the Caucasus molt commences 
a month or so later than in the lowlands (Radde, 1889). The young 
generally molt slightly later than adults. 

The summer hair coat consists of uniform guard hair which is 
sparser than in winter. Growth commences from August when 
underfur is also seen. In October the rust-colored summer coloration 
alters to a dark brownish-gray. After this the guard hair develops 
only insignificantly, lengthening up to December when it is longest 
and already duller than in autumn. When autumn is cold the winter 
coat develops earlier. 

At the age of 10 years teeth start falling out and females become 
barren. The maximum age of females is usually 11 or 12 and of 
males about 16 years (Kalnin'sh, 1950). 

Enemies, diseases, parasites, mortality, com.petitors, and popula- 
tion dyiiamics. Over much of the range the wolf is one of the main 
212 enemies of roe deer. This predator subsists at some places mainly on 
roe deer, as a result of which its seasonal distribution and move- 
ments are often determined by the corresponding phenomenon in 
the life of roe deer (Podarevskii, 1936; and others). In Belovezh 
Forest, which is rich in some species of ungulates, remains of roe 
deer constituted 18.4% of wolf food (in over 1,100 samples of feces 
and stomach contents collected from 1946 to 1950). Wolves wreak 
maximum damage in winter. For example, in the winter of 1946 to 
1947 when the snow height in Belovezh Forest was 55 cm, roe deer 
remains were seen in 28.2% of the food of wolves (Gavrin and 
Donaurov, 1954). In 11 'men preserve roe deer are the main species of 
ungulates and their remains were found in 95.9% of all the data on 
food (395 samples) of wolves (in the summer 84.7%, winter 99.2%, 
and spring 100%). In this preserve 10 to 12% of the roe deer popula- 
tion are killed annually by wolves (Averin, 1949). Wolves cause 
exceptionally great damage to roe deer in Bashkir (Kirikov, 1952) 
and Irkutsk districts (Fetisov, 1953), Yakutia (Belyk, 1948), Primor'e 
territory (Abramov, 1954), and elsewhere. 

Compared to wolf, all other predators are less important except, 
perhaps, for lynx (yellow-throated marten in Primor'e territory and 
red fox for European roe deer). In Altai preserve roe deer remains 
constituted 58.9% of the number of all quarries of lynx (56 observa- 
tions on the tracks of lynx; Dul'keit, 1953). Thus roe deer represent 
the main food source of lynx in the Altai. Similar findings with 
respect to lynx and roe deer have been recorded in the central Urals 
(Sabaneev, 1875) and the Caucasus (Dinnik, 1910; and others). How- 
ever, roe deer remains were found only in 3.6% of the data (136 



295 

samples) on feces of lynx from Caucasian preserve (Kotov, 1958). In 
the feces of lynx from Belovezh Forest the remains of roe deer were 
found in 20% of the total available data (Yurgenson, 1955). In some 
regions (Urals and Altai) lynx follow close behind roe deer during 
their migration to wintering sites. 

Red fox often attack young roe deer, especially in the first few 
weeks after birth. However, in Crimea even adult roes, mainly ema- 
ciated and sick females, are also attacked frequently. From 1937 to 

1940, when the fox population in Crimean preserve was very high, 
about 200 roe deer were killed annually by them. In those years over 
50% of all established cases of death of roe deer were due to attacks by 
fox (Rukhlyadev, 1948). Epizootic diseases decimated the fox 
population of Crimea in 1946 and 1947. Yet in 1948 and 1949 fox 
were responsible for 36.8% roe deer mortality (P. A. Yanushko). Very 
large numbers of roe deer have been killed by fox in the Crimea in 
snowy winters (Dal', 1930; and others). In Germany the fox is 
regarded as the main enemy of roe deer; the maximum number of 
adult animals are killed by fox in winter when the snow cover is 
thick and crusts have formed (Raesfeld, 1956). In the Urals, the 
Caucasus, and especially in Siberia, where the large form of roe deer 
lives, fox almost exclusively attack the animals in their first year. 

Many other predators, including some birds, can be considered 
enemies of roe deer but their role in population regulation is not 
significant. Such predators are bear (more dangerous for young 
deer), wolverine, tiger, leopard, red wolf, jungle cat [F. chaus], wild 
cat [F. silvestris], marten, golden eagle, eagle-owl,'^" and others, 
which threaten the life of young and juvenile roe deer. Wandering 
wild dogs take quite a large toll of roe deer; for example, in April, 

1941, at the end of a snowy winter in Bashkir preserve 17 deaths of 
roe deer caused by wandering dogs were recorded (Kirikov, 1952). 

In severe and snowy winters, and also in winters with prolonged 
ice crusts, not only are a large number of roe deer killed by wolves 
and other predators, but many die due to emaciation, diseases, '^' etc. 
Instances of mass mortality of roe deer in such winters are known in 
213 much of the range and have been described time and again in the 
Crimea, the Caucasus, the Urals, southern Siberia, etc. (Nasimovich, 
1955). In Latvia, in the severe winter of 1939-40 about 20% of all roe 
deer died (Severtsov, 1941). After the snowy winters of 1945 to 1947 
roe deer were totally decimated at some places in the central Urals 

Remains of roe deer fawns were found in nests (Fetisov, 1953). 
Especially inflammation of the lungs occurring after prolonged cfiases by 
predators and tfie subsequent intake of snow by the flushed animal (Ushkov, 1954). 



296 

and reappeared only after some years. In the steppe regions roe deer 
are threatened by glaze ice which facilitates wolf chases of ungulates. 
A large number of roe deer died in the winter of 1948-49 in central 
and southern Kazakhstan when ice crusted on the heavy snow cover 
(Sludskii, 1953). In the mountains of western Caucasus death of roe 
deer due to snow avalanches has been reported (Nasimovich, 1955). 

Roe deer are exposed to several infectious diseases: foot-and- 
mouth disease, cattle plague, Siberian ulcers, necrobacillosis, erysi- 
pelas, salmonellosis, tuberculosis, pasteurellosis, pyroplasmosis, and 
others. Some were detected in Crimean preserve (Rukhlyadev, 1939 
and 1948), in the Caucasus (foot-and-mouth disease; Dinnik, 1910), 
and Moscow Zoological Garden (Sludskii, 1954; Fromkhol'd, 1958). 
Instances are known of mass mortality of roe deer due to epizootic 
diseases, the reasons for which are yet to be explained: in 1910 and 
1912 in Belorussia (Serzhanin, 1953), 1926 in Azerbaidzhan (Lavrov, 
1929), 1930 in Dzhungarsk Alatau (Shnitnikov, 1936), etc. 

Eighty species of helminths have been identified in European roe 




fc.**.— *— лЭ^ -J*- *- 



Fig. 65. Track of roe deer climbing a mountain. Middle course of Kan River. 
Irbeiskoe region of Krasnoyarsk territory. March, 1952. Photograph by K.T. 

Yurlov. 



29-7 

deer alone; of these, 68 have been recorded in the USSR (Belyaeva, 
1958). In the Crimea moneziasis of the intestines and dictyocaulosis 
of the lungs are particularly dangerous, sometimes causing roe deer 
mortality (Rukhlyadev, 1948). Many roe deer are severely infected 
with different varieties of Trichostrongylus, which attack the respi- 
ratory tract and in zoological gardens often lead to animal mortality. 
Cases of coenurosis have also been recorded. Roe deer of Aksa- 
Dzhabaglinsk preserve were once severely infected with Skrjabinema 
ovis (Boev et al., 1957). In Crimean preserve 12 cases were confirmed 
of roe deer mortality due to theilerosis, a disease caused by blood 
parasites (Rukhlyadev, 1948). 

Nasopharyngeal {CepheJiomyia stimulator and Pharyngomyia 
picta) and dermal {Hypoderma capreoli, H. diana, and others) 
gadfly larvae parasitize roe deer. Larvae of C. stimulator sometimes 
penetrate the trachea and brain of roe deer. In Germany such events 
occurring in the period when the larvae of С stimulator attain large 
size (April-May) may account for 55% of the annual mortality of this 
species (Grunin, 1957). Among other parasites of roe deer are the 
blood-sucking Lipoptena cervi, hair louse Trichodectes tibalis, and 
itch mite Sarcoptes (Acarus) scabei. Roe deer also suffer from horse- 
flies, mosquitoes, gnats, ticks, etc. 

The most serious competitor of roe deer among the ungulates is 
the red deer. The adverse influence of the latter on the population of 
roe deer (destruction of winter food resources) has been observed in 
Belovezh Forest, Altai, and Crimean preserves, but as yet not ade- 
quately studied. Wild boar used to destroy large numbers of roe deer 
fawns in the Crimea (Dal', 1930). 

A large number of young roe deer evidently die in the first few 
weeks or months after birth. In summer months the young consti- 
tute 26.9% of the total roe deer population in Crimean preserve; by 
the end of winter they comprise only 15.4% (P. A. Yanushko). In 
Belovezh Forest, depending on the severity of winter and the wolf 
population, 49 to 80% of roe deer young have perished by spring in 
various years (Sablina, 1955). Judging from long-term observations 
in Il'men, Crimean, and Belovezh Forest preserves, females predomi- 
nate in roe deer populations and account for 54 to 68% of adult 
animals. At birth the sex ratio is nearly 1:1 or even slightly in favor 
of males. Thus the mortality of males is somewhat more than that of 
females and evidently the result of death during combat, emaciation 
during rut, and so on. 

In Gatchina (Leningrad district), where 59 Siberian roe deer were 
released in 1896, the herd growth over a 12-year period averaged 26%. 
In some game farms of western Europe the herd growth index is 



298 

much higher (Sever tsov, 1940 and 1941). 

Field characteristics. Roe deer are of relatively low stature (height 
at withers usually less than 90 cm versus 130 to 160 cm in red deer) 
and have a relatively short trunk and small, almost erect antlers with 
a limited number (up to three) of tines. These features make identifi- 
cation of roe deer among other members of the deer family quite 
easy. In summer the main coloration is rusty, much lighter on the 
lower parts, and the speculum around the tail poorly defined. In 
winter the general coloration is gray or grayish-brown, sometimes 
grayish-rusty, gradually darkening toward the rear and turning 
lighter downward, and the light-colored speculum around the tail is 
distinct. Young are spotted in the latter half of summer. Females are 
only slightly smaller than males; in winter, when males have no 
antlers, their differentiation from females at a distance is quite diffi- 
cult. The neck in males is very short and thick and the hair hangs 
down under the abdomen (Kalnin'sh, 1950). 

Roe deer hoofprints are small, blunt, and rounded in males and 
relatively long and narrow in females. When moving slowly the 
distance from the center of one print to that of another averages 45 
cm in adult Crimean roe deer; they usually jump 1.5 to 2.0 m but 
when very frightened even up to 7.5 m (Dal', 1930). 

In winter roe deer try out several places (up to six) before select- 
ing a bed. A bed in snow is very small and not equal to the body size 
since the legs are tucked in. In regions with severe winters roe deer 
usually lie on the ground and leaves after scraping away the snow 
first; they often lie directly on dry snow, however. In winter males 
are readily distinguished from females by the position of the brown 
urine patch on the track. 

In winter feces are in the form of longish pellets (10 to 14 mm x6 




Fig. 66. Winter feces of roe deer (magnified 1.5 times). Primor'e 
territory (from Formozov, 1952). 



299 

to 9 mm) and in spring formless cakes; both types of feces are seen in 
summer. Tfie pellets are much smaller (0.7 mm x 0.4 mm) for six- 
month-old deer. Identification of sex from the form and size of feces 
is unreliable. 
215 In the period of rut roe deer stamp out characteristic circles 
around trees, beat the turf, batter branches of shrubs with their 
antlers, etc. (see above). Unlike red deer and elk, roe deer never touch 
bark. Like other ungulates, they often make trails, especially near 
salt licks, etc. 

Roe deer emit a prolonged squeak and a short, interrupted, very 
loud, and hoarse bellow ("gau, gau"); females bellov/ very rarely and 
their voice is less coarse. Roe deer call most often when frightened. 
The young emit a high and prolonged squeak. 

In winter roe deer seldom, if ever, bark. (A.N.) 

Economic Importance 

Throughout most of the nineteenth century, and at some places in 
the first two decades of the twentieth, roe deer were caught in the 
Soviet Union in very large numbers. Hunting roe deer was particu- 
larly practiced in southern Siberia, the Far East, and the Urals, 
acquiring commercial proportions. The largest number of roe deer 
were caught in winters of heavy snow. For example, in the exces- 
sively snowy winter of 1877 78 about 150,000 roe deer were caught in 
the Amur region (Turkin and Satunin, 1902). Mass shoots were 
carried out in autumn and early winter at places of migration of roe 
deer on ice and while swimming across the Amur, Ussuri, and to a 
- lesser extent also across the Yenisey, Abakan, Bir, Irtysh, and other 
rivers. In the Amur hunters following "moving goats" [roe] for 200 
to 300 km; in successful years a hunter bagged 50 to 100 deer (Nasi- 
movich, 1955). 

Various methods of mass capture of animals were practiced in 
different parts of the range: nooses and traps set in passages of 
specially erected fences, chasing in the period of migration, through 
deep snow, on ice, etc. Roe deer were caught not only in the cold 
season of the year, but also in summer — by voice imitation (luring 
females by imitating the voice of the young), at salt licks, in trails 
etc. The meat of roe deer represented an important food source in 
many towns of Siberia. 

Thoughtless and often savage hunting adversely affected the 

"^ In the winter of 1887 to 1888 the carcass of a roe deer in the Blagoveshchensk 
market cost 1.60 to 1.70 rubles. 



300 

reserves of the species and the population of roe deer in most regions 
was depleted. For this reason after the October Revolution hunting 
was controlled in the Soviet Union; roe deer shoots were greatly 
restricted and at places totally prohibited (the Ukraine, Belorussia, 
and several regions of the Russian Soviet Federal Socialist Repub- 
lic). In recent years maximum numbers of roe deer have been caught 
in southern Siberia'^^ and Primor'e territory, reaching 40,000 to 
50,000 animals per year, followed by Kazakhstan and Kirgizia (up to 
5,000 to 10,000), the Urals, etc. In the Caucasus, Crimea, and Baltic 
region hunting is purely a sport, often limited only to males and 
that too on a very small scale, and at places not permitted every year. 
In the Asiatic part of the USSR the reserves of roe deer in general are 
quite considerable (see above) and hence a catch of 50,000 to 60,000 
animals per annum is permissible. The most rational periods for 
hunting are October and November and from November onwards in 
the southern parts of the country. 

At present the most prevalent methods of hunting are: shooting 
from ambush, catching by use of pens, waiting at places of regular 
visitations and feeding grounds, chasing on sledges (in Siberia), etc. 
In spite of legislation prohibiting it, at places, for example the Altai, 
roe deer are increasingly caught in summer with mechanical traps or 
chased in winter (the Urals) on heavy snow and ice crust. Dogs are 
usually employed only for locating wounded animals. In the past 
hunting with hounds was often practiced in the Caucasus, the Urals, 
and elsewhere. 

The net weight of meat from a southern Russian roe deer is 
usually not more than 14 kg and from a Caucasian an average of 15 
kg, while Siberian and Middle Asiatic animals yield slightly over 20 
kg (very rarely up to 30 kg), representing roughly 60% of live weight. 
Depending on size and fattening a roe deer can provide 1.5 to 2.5 kg 
fat. The meat of roe deer is less valued than that of other species of 
wild ungulates. 
216 Roe deer hide is utilized for articles of apparel and chamois. Late 
September and October hide is considered the best and called "Bar- 
low hide"; by December the hair has become less stable and the hide 
poor in quality. Hides are mainly used in fur coats (with the hair 
outside), caps, and more rarely other articles such as flying boots, 
mittens, sleeping bags, and cloaks. In the past hides were also con- 
verted into collars, jackets, breeches, belts, and saddle cloths; 
summer hides were used in small rugs, and so on. Fur coats are 

In 1924 about 47,000 roe deer were caught in Tuva Autonomous District alone 
(Yanushevich, 1952). 



301 

warm but shortlived since the hair falls out rapidly; properly cared 
for a coat may last three to five years. Antlers with the frontal bone 
serve as wall decorations (in the nineteenth and early twentieth cen- 
tury they were exported to Germany) and are also converted into 
small articles. 

In cultivated areas roe deer inflict damage only when they occur 
in large numbers. Then they trample and browse tender saplings, 
damage winter crops (rye and others), and spring oats. In summer 
they damage cereal crops (in which they make beds), clover fields, 
and vegetable patches (trampling and eating potato tops); in winter 
they stamp hay left over in fields. In Belovezh Forest roe deer some- 
times threaten self-sown pine (browse them in winter), oak, beech, 
and maple, but the damage by and large is insignificant (Sablina, 
1955). In Il'men preserve aspen seedlings are most intensely dam- 
aged (100%), followed by pine and birch (40 to 50%). When 10- to 
15-year-old pine and birch trees grow beyond the reach of roe deer 
they usually recover. Hence the damage to forest trees caused by roe 
deer is also insignificant (Averin, 1949). In Estonia roe deer damage 
most intensely aspen forests (heartrot extends from the point of 
damage on the trunk). In Latvia two- to three-year-old pines suffer 
and at places where roe deer are numerous even eight-year-old pines 
are not spared (Kalnin'sh, 1950). Judging from the experience gained 
in Czechoslovakia, the negligible damage caused by roe deer is more 
than compensated by the value of this species as a game animal. 

Roe deer have been released time and again into private farms for 
purposes of raising them as game animals. In 1891 eleven Siberian 
roe deer were imported into Belovezh Forest where they acclimat- 
ized and reproduced (Kartsev, 1903). The 59 animals released in Gat- 
china game farm (now in Leningrad district) in 1896 had multiplied 
to 1,068 animals by 1909 (Severtsov, 1940), but later became extinct 
there. Right up to World War I, 15 to 20 European roe deer were 
held at Ramon' (Voronezh district) in a pen (free roe deer there had 
disappeared). Later they were brought into Usmansk Forest where a 
few were conserved; they are now slightly greater in numbers 
(Nikul'tsev and Predtechenskii, 1957). 

Commencing with 1925 small batches of roe deer from the Cri- 
mea and other places have been released in the Ukraine (Diken'ka, 
Pechenegi, and other regions); almost everywhere these roe deer were 
later killed by wolves, poachers, etc. Ten roe deer were brought from 
Kirgizia to Mordov preserve in 1940. After 1943 they were no longer 
sighted there [Letopis' prirody Mordovskogo zapovednika (History 

Local roe deer were also present in the forest. ^ 



302 

of Mordov Preserve)]. In 1932 and 1939 a total of 12 roe deer were 
brought into Zavidov game farm (north of Moscow) from the south- 
ern Urals. By 1941 they numbered 50 to 60 animals and by 1945, after 
the War, about 40 (Danilov, 1947). Roe deer are still seen in this 
farm. European and Siberian roe deer have been released time and 
again around Moscow on Losin Island. In 1941 there were about 80 
animals — all killed during the War (Formozov, 1947). In 1950 and 
1952 some were released in Sobin and Petushin areas of the Vladimir 
district. These animals survived, reproduced, and later spread into 
adjoining regions of the district (Sysoev, 1955). In 1954 ten Siberian 
roe deer were released in Pereslav game farm (Yaroslav district), and 
in 1957 forty (from Primor'e territory) in Dubna (Moscow district). 
Roe deer in recent years have been released in other regions of the 
Soviet Union also. 

In nature roe deer are pugnacious, often attack man, and suffer 
from various diseases. (A.N.) 

279 GENUS OF ELK* 

Genus A Ices Gray, 1821 

mb.Alce. Frisch. Natur-Syst. d. vierjiiss. Thiere, p. 3. 
\^2\.Alces. Gray. London. Med. Reports, vol. 15, p. 307. Cervus 

alces Linnaeus. 
ISil. A Ice lap hus. Gloger. Handbuch d. Naturgesch., p. 143. Substi- 
tute for Alces. 
1902. Paralces. J. Allen. Bull. Am. Mus. Nat. Hist., vol. 16, p. 160. 
Substitute for Alces. (V.H.) 

Alces are telometacarpal deer of large size (the largest forms of 
the subfamily and family); heavy build with a relatively short trunk; 
high withers; large, massive, heavy, and highly elongated head; and 
long legs. 

Hooves of middle toes narrow, long, and sharp; lateral hooves 
large and set quite low. 

Skull relatively large, highly elongated (width in region of zygo- 
matic arch slightly over one-third its total length), with a well- 
developed facial part: distance from center of orbit to anterior tip of 
premaxillae roughly double the distance from center of orbit to rear 

•Termed "moose" in North America, where the name "elk" is often apphed to the 
wapiti (Cervus elaphus). 

'in view of some confusion associated with the name A Ice (often apphed to giant 
deer), the International Nomenclature Committee has recommended the use of Gray's 
name. 

Functional, and facilitate movement on soft marshy soil and on snow. 



303 

level of occipital condyles (distance from anterior molar to end of 
premaxilla roughly 1.5 times upper tooth row). Nasal processes of 
premaxillae do not reach nasals. Latter very short, roughly one-third 
shorter than upper tooth row; nasal passage very large and markedly 
longer than upper tooth row. Preorbital depression on lacrimal 
bone well developed but small and set away from rim of orbit; 
ethmoid pit distinct. Rear of vomer low and does not divide poste- 
rior openings of choanae into two parts. Tympanic bullae very 
small, do not project beyond glenoid fossae, and covered with sharp 
bony projections; auditory meatus long. Bony pedicels of frontals set 
straight on sides (perpendicular to sagittal plane of skull) and fall in 
plane of frontal surface of skull (the only instance in the family). 
Lower jaw highly elongated. Height of symphysis nearly equal to 
length of tooth row. 

Upper canines absent in males as well as females and their 
sockets closed (rudimentary canine a rare exception). Molars rela- 
tively large, massive, and broad with low crowns. Lateral walls of 
teeth incline inward, as a result of which masticatory surface is 
smaller than tooth base. Incisors and lower canines relatively 
poorly specialized and differ little from each other in size and form. 
Dental formula: 

.00 3 3 „^ 

1 — , с — , pm — , m — =32. 
3 13 3 

Only males sport antlers, which run straight out to the sides, 
quite horizontally, slightly turning upward near the end, and gener- 
ally set sideways. Antlers either wholly (except for small section of 
beam at base) webbed with small tines set forward, sideways, and 
218 backward, or, along with palmate portion exhibit a special tine 
quite separated from it (sometimes wholly and in others not so); this 
special tine is set forward and usually bifurcate at the end, or pal- 
mate portion not developed at all and antlers branched like those of 
"deer"; antler beam circular in section. Antler surface, especially of 
beam, uneven and rough; color deep brown. 

"' In form and relative size molars of elk have common features with those of 
giraffe, evidently due to their feeding on similar types of food (branches). 

"^ The homology of parts of elk antlers is difficult. The anterior tine of the 
corresponding part of the palmate portion is taken as representing a fused brow and 
middle, or second and middle, or only middle tine of a typical deer antler. Usually the 
brow tine is assumed to be absent in elk. The palmate portion is formed by the 
flattened end of the beam or the rear tines fusing together but set forward or sideways 
(as a result of a twist in the beam) (i.e., the palmate portion is homologous to the rear 
tines of reindeer and fallow deer). Thus, the rearmost inner tine (set toward the neck) 
would form the beam end and the tine in front of it would be the rear one. 



304 

Ears very long and broad. Tail short. Muzzle, i.e., upper lip and 
nasal region, very large, broad, and viewed from the front, appears 
cut in straight line ("squarish"); overhangs lower lip and has a 
complex inner cartilaginous skeleton. Muzzle covered with hair 
except for tiny triangular or rhomboidal patch situated midway on 
front lower surface of upper lip. 

Dewlap, usually more prominent in males present on throat. 

Preorbital glands well developed but not very large. Hoof glands 
present only on hind limbs and resemble saccate invaginations with 
a small opening on the anterior surface. Tarsal and metatarsal 
glands small; latter set high (at the level of calcaneal joint) on the 
inside of legs. 

Coloration uniformly dark (blackish-brown or almost black), 
legs considerably lighter than trunk, and speculum on rear of thighs 
absent. Hair on neck and withers elongated. Coloration of young 
uniform (without light spots). Female somewhat smaller than male. 
Teats four. 

These are inhabitants of taiga, mountain taiga, and mixed 
forests, with a preference for lakes and marshy regions, in part 
forest-tundras and forest-steppes, and bottomland deciduous forests 
in steppes. Evidently, unlike most other members of the family, they 
are monogamous. 

The range of the genus is associated with the forest of the north- 
ern hemisphere and is divided into two parts — American and Asian. 

In North America the range (reconstructed) occupied much of 
the forest zone (apart from purely deciduous forests). Its northern 
boundary in the northwest covered the Yukon basin, ran to the 
Mackenzie estuary, and descended from there in an irregular line to 
the southwest corner of Hudson Bay, and in the east to the St. 
Lawrence Gulf. Farther away the range encompassed all of Nova 
Scotia and New England (elk have been acclimatized in Newfound- 
land) and formed a prominent extension southward along the 
northern parts of the Allegheny mountains. More to the west the 
southern boundary ran through the region of Lake Superior 
(between upper [Lake] Michigan and Huron), and encompassed the 
northern parts of Wisconsin, Nebraska, and North Dakota; passing 
north of [eastern] Montana and southern Saskatchewan, it des- 
cended in the form of a promintory along the mountains up to the 
Yellowstone (around 45° N lat.). From there, gradually coming 
closer to the Pacific Ocean coast, the boundary departed northward 
into Alaska. 

In Eurasia the range of elk (reconstructed) occupied the entire 
forest zone of Europe, in the west to the Rhine estuary and to the 



305 

Vosges; in the south up to the northern foothills of the Alps and to 
the Carpathians, part of the steppe zone (at least the Don basin and 
probably the Volga), the Caucasus and western Trans-Caucasus, all 
of the forest zone of Siberia including forest-tundia (at places even 
tundra) and forest-steppe, fofest parts of Mongolia (Khangai and 
Pri-Hobsogol), forest regions of the Far East (except Kamchatka), 
the Ussuri region, and northeast China. '^^ On the whole the range is 
extremely typical of circumboreal ranges of forest (mostly taiga) 
forms. 
220 At several places the range of elk has shrunk and continues to 
diminish (America); simultaneously a perceptible enlargement of it 
has occurred in the USSR, mostly in the south and partly in the 
north. 

The genus A Ices occupies an extremely isolated position among 
present-day telometacarpal deer (Odocoileinae) and not one of the 
other genera of the subfamily reveals any close relationship to it. 
Some researchers have isolated this genus in a special tribe. 

The origin of the genus cannot be explained with any degree of 
accuracy. A wholly typical form of it is seen in the Early Quaternary 
(Gunz) period. The genus Aires should evidently be derived from 
Pliocene Eucladocerus and its origin placed in the Upper Pliocene. 
The North American Quaternary Cervalces is close to Alces; the 
former in some respects can be considered intermediate between the 
two genera Alces and Eucladocerus (Flerov, 1950). Eucladocerus 
prospered in the Middle and Upper Pliocene and partly in the Early 
Quaternary period of Europe and Asia. Evidently elk, in spite of 
their affiliation to the predominantly American form of deer, should 
be considered as having originated in Europe. ^"^^ 

Some species of Pleistocene elk have been described but most of 
these descriptions are baseless. At least in the USSR only one inde- 
pendent Quaternary species occurred, the broad-fronted elk Alces 
latifrons Dawk., typical of the Lower Pleistocene. Its descendants 
survived in the Upper Quaternary period and by then bore resemb- 
lance to present-day species (Gromov, 1948). They were distributed 
somewhat more extensively than in the historic period and encoun- 

For datails, see description of range of elk given below. 

According to some paleontologists (Simpson, 1945) genus Eucladocerus is 
neither related to genus Alces nor even subfamily Cervinae; it has affinities with the 
giant deer (Megaloceros). Accordingly, only the Pleistocene North American Cerval- 
ces is associated with Alces as the most proximate and ancestral form. The origin of 
the genus Alces is thereby also shifted to American territory in which most other 
telometacarpal deer (Odocoileinae) exist and prosper. Giant peat deer, having huge 
antlers with palmate or webbed portions and externally similar in general to elk, 
according to a view commonly held, are not related to genus Alces either. 



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307 

tered (often even in the Neolithic period) in the steppe regions of the 
Ukraine, lower Volga, Trans-Caucasus (Rion basin), on the Black 
Sea coast of the Caucasus, and in England and Ireland. In western 
and central Europe, however, elk did not penetrate south of the 
Pyrenees and were not found in the Balkan and Apennine 
Peninsulas. 

Until recently the genus purportedly included four, three, or two 
species. Some contemporary American researchers still accept two 
species of elk, i.e., Eurasian and American. Yet it is absolutely clear 
that the genus consists of only one species, Alces alces Linn, (about 
5% of the species of the subfamily). 

These animals represent hunting and table animals and are 
sought for their meat and hide. 

There is only one species of elk in the USSR, Alces alces Lin- 
naeus, 1758 (0.3% of species of Russian fauna). 

Elk are taiga and mountain taiga animals with a preference for 
swampy areas, partly occupying mixed forests, and more rarely 
forest-steppes, forest-tundra, and tundra. 

Elk as hunting and table animals are very important at places 
(western Siberia) to the local populace. Hunting of elk is prohibited 
in many parts of the country or allowed only by special pemiission 
(license). (V.H.) 

ELK, SOKHATYI^°' 

6. Alces alces Linnaeus, 1758 

222 lib's. Cervus alces. Linnaeus. Syst. Naturae, 10th ed., vol. 1, p. 66. 
Sweden. 

ViSb.Cervus alee. Boddaert. Elench. animalium, vol. 1, p. 135. 

\%22.Ceruus americanus. Clinton. Letters Nat. Hist, and Intern. 
Resources State New York, p. 193. New York. 

\W2.1 .Cervus coronatus. Lesson. Man. de Mammal., p. 356. 

\ЪЪО. Alces europaeus. Burnett. Quart. Journ. Sc. Lit. Art., 1829, 2, p. 
353. Nomen nudum. 

ISbl.Alces machlis. Ogilby. Proc. Zool. Soc. London, 1836, p. 135. 
Substitute for alces L. 

1842. Alces antiquorum. Ruppel. Mus. Senkenb., vol. 3, p. 183. Sub- 
stitute for alces L. 

^'" Male = los' [elk] bull; female = cow, losikha. Attempts (Flerov, 1935 and 1952) to 
intioduce into the Russian language the popular American name "moose" for east 
Siberian elk should be vigorously opposed since there is no justification whatsoever 
for such usage. 



308 

IS^^ -Alces palmatus. Gray. List Mamm.. Brit. Mus., p. 142. 
\^60.Alces jubata. Fitzinger. Naf. Sdugeth., vol. 4, p. 86. Substitute 

for alces L. 
\S&1 .Alces cameloides. Milne-Edwards. Ann. Sc. Nat. ZooL, vol. 7, 

p. 377. Manchuria. 
\S99. Alces gigas. Miller. Proc. Zool.Soc, Washington, vol. 13, p. 57. 

Kenai Peninsula, Alaska. 
1902. Alces bedfordiae. Lydekker. Proc. Zool. Soc, London, 1902, 

vol. 1, p. 109. Eastern (?) Siberia. 
1910. Alces machlis typicus. Ward. Rec. Big. Game, 6th ed., p. 99. 

1910. Alces pfizenmayeri. Zukowsky. Wild undHund, vol. 16, p. 807. 
Aldan River, Yakutia. 

1911. Alces machlis yakutskensis. Millais. The Field, vol. 118, p. 113. 
Aldan River, Yakutia (name based on same material as in 
preceding reference). 

19\3. Alces machlis uralensis. Matschie. Veroff. Institute J agdkunde 
Neudamm, vol. 2, p. 155. Southern Urals. 

191S. Alces machlis meridionalis. Matschie. Veroff. Institute J agd- 
kunde Neudamm, vol. 2, p. 156. Samar province. 

\9\Ъ. Alces machlis tymensis. Zukowsky. /4гс/г. /. Naturgesch, vol. 
80A, no. 9, p. 42. Western Siberia, Тут River. 

1915. Alces machlis angusticephalus. Zukowsky. Archiv. f. Natur- 
gesch, vol. 80A, no. 9, p. 44. Yenisey province. 

\92Ъ. Alces alces bedfordi. Bobrinskii. Opredelitel' okhotnich'ikh i 
promyslovykh zverei (Identification of Game and Table Ani- 
mals). Ussuri region. 

1955. Alces alces caucasicus. Vereshchagin. Zoo/ogzc/i. Zhurnal, vol. 
34, no. 2, p. 461. Urukh gorge, central part of northern slope of 
Great Caucasus. (V.H.) 



Diagnosis 

The genus Alces contains only one species. 

^"^ The author of this work attempted to identify precisely the typical habitat of 
the form described by Lydekker (1902) since it was shown in an extremely generalized 
manner. N.A. Bobrinskii mistakenly used the patronymic name given initially to the 
female form with a masculine ending. This error was repeated later by the author 
(1944) and by others. There is no basis for Bobrinskii giving this name to Ussuri elk, 
as assumed by Buturlin (1934), who recognized neither the slip nor error made by this 
author. In general this name and its given outline, initially accepted by Lydekker 
(1912), is no more than a synonym since Manchurian-Ussurian elk ought actually to 
be labeled cameloides, the name assigned to them in 1867 by Milne-Edwards. 



309 

Description 

Large deer, the largest living species of the family. In general 
appearance very massive and powerful, somewhat heavy but well 
built. Build differs markedly from other deer, primarily in long legs, 
powerful and massive chest and shoulders, and large and heavy 
head. Neck and head usually held low and fairly horizontal. 

Since the legs of elk are very long the trunk, though of normal 
length, appears relatively short and the withers high; the latter with 
223 its long coat of hair forms a humplike structure. Back straight, 
sacrum slightly lower than withers, croup relatively poor and slop- 
ing, tail very short, much less than one-half length of ear and not 
discernible in an active' animal. Neck relatively short, thick, and 
massive. Head relatively very large, long (roughly equal to neck in 
length), narrow, and aquiline. Upper lip very large and massive, as 
though inflated, squarish when viewed from below, and proini- 
nently overhangs lower lip. Nostrils large and set low. Ears very 
large and broad, generally oval in form, but pointed at tips and 
highly mobile. Eyes relatively small. Small preorbital glands pres- 
ent. Dewlap fairly long with a triangular profile but sometimes 
sausage-shaped; its size maximum in animals three to four years old, 
becoming shorter and broader later. Dewlap length reaches 35 to 40 
cm, usually 20 to 25 cm. 

blooves large, long, and narrow; strongly pointed forward. In 
females somewhat narrower and sharper than in males. Lateral 
hooves relatively large, set low, and when moving on soft soil rest on 
the ground and carry part of the body weight. 

In general appearance cows are indistinguishable from bulls, but 
body build of former somewhat more slender and chest, shoulders, 
and withers less massive. In the first few months particularly sharp 
differences between trunk and limbs apparent in both sexes — a dis- 
proportionately small trunk on particularly long legs. Moreover 
their head is relatively small and the upper lip not swollen but nearly 
normal. In their first winter the body proportions of young elk stand 
out even more prominently compared to those of older animals; 
fairly typical proportions of adult animals are not attained earlier 
than in the second year. 

Antlers of elk are extremely variable in size and structure, more 
so than those of other Russian deer except, perhaps, reindeer. Age 
variability, regional variability, and sometimes extreme individual 

^"^ Elk, unlike other deer but like the giraffe, amble because of their long legs. 
See characteristics detailed under genus. 




221 



Fig. 68. Elk (A Ices alces L.). Sketch by A.N. Komarov. 



311 

variability are apparent. Typical elk antlers consist of a short beam 
set off from the skull horizontally and perpendicular to the sagittal 
plane of the body. The plane of the flattened and fairly curved 
palmate portion is directed forward to a lesser extent, more sideways, 
but mainly backward. When the head is held in a horizontal posi- 
tion, the plane of the palmate portion is almost horizontal, rising 
slightly only in the rear. Tines are directed from the palmate portion 
forward, outward, and backward, but not inward (not toward the 
neck). They are also set slightly upward in continuation of the 
curved shape of the palmate portion. All the tines are more or less 
identical and form a frame around the periphery of the palmate 
portion uniformly, but more often the tine directed forward is larger 
than the rest and somewhat isolated from the palmate portion. 
Often, individual tines are quite independent even on the other parts 
of the palmate portion, but more often in the posterior or postero- 
lateral parts; an appreciable indentation occurs in the palmate part. 

Antlers of this type may attain very large proportions. However, 
in most cases the flat zone of the palmate portion itself is small and 
the tines long. Usually, the larger the palmate part the shorter the 
tines and vice versa. Maximum-sized antlers have an extremely long 
and broad (up to 60 cm or more) palmate section with short tines, 
often with a prominent forward tine (on the palmate section) which 
bi- or trifurcates at its tip. 

The second antler type is also palmate but a massive separate tine 
stands prominently forward and is usually bifurcate at the tip, 
224 resembling somewhat two forward tines from the anterior part of the 
palmate section. Several intermediate varieties occur between this 
and the typical antler form. Both types are similar and differences 
only quantitative. 

The third antler type — the "cervine" type antler, completely 
lacks a palmate section and the beam is fairly short and thick, sym- 
metrically branching in a fairly horizontal plane. The tines are set 
forward, sideways, and backward, and upcurved. 

Between the antler types described above, especially the first two, 
several transitional forms occur, each exhibiting wide variations. 
Antlers with a small palmate portion and long tines, usually with a 
massive bifurcate tine prominently set forward, are most common 
among Russian elk. The number of tines on particularly large 
antlers can reach 36 (on a pair of antlers) and probably even more. 
Some geographic similarities are seen in these types, especially in 
"cervine" type antlers (see "Geographic Variability"). 

Age-related changes in antlers are as follows. A two-year-old bull 
carries a small unbranched stump on which two tines (forks) form in 



312 




224 Fig. 69. Some types of antlers in elk (A. alces). 

1— Well-developed palmate antlers of central Russian elk {A. a. alces); 2— large antler 
of elk from Bryansk region (maximum length 68 cm); 3 — typical antlers of elk {A. a. 
alces) from western Siberia (Dem'yanka River); 4 — unusually large antlers of western 
Siberian elk with long and broad palmate portion (Dem'yanka River); 5 — typical 
antlers of Ussuri elk(.4. a. cameloides). From Kaplanov (1948) and Flerov (1952), with- 
some modifications; antler of Bryansk elk from A.V. Fedosov. 



the third year. Later, the antler changes in no regular or fixed pat- 
tern except that generally the older the animal the larger and more 
massive its antlers, the larger the palmate portion, and the shorter 
the tines. In extreme old age deterioration and secondary simplifica- 
tion of antlers set in which, evidently, can be extensive." 
225 The hair of elk is similar to other deer — coarse, slightly wavy and 
thick, with air cavities (especially in winter pelage), and brittle. 
Winter hair on the trunk is about 10 cm or more in length and even 
slightly longer on the back. Hair is particularly long on the withers 
(about 16 to 20 cm) and on the upper part of the neck where it forms 
a characteristic mane. Long hair on the withers is largely responsi- 
ble for the "hump" so characteristic of the general shape of elk. On 

The structure, development, variability, and other features of elk antlers have 
hardly been studied. This is particularly true of Russian elk. 



313 

the sides of the neck the hair is not so long, only slightly longer than 
that on the body sides. 

The head is covered with short and slightly softer hair, which is 
particularly short but very resilient on the entire anterior part of the 
muzzle. Only at the center of the upper lip does a minute bare spot 
occur, which is oval, pyriform, or triangular in shape. It is so small 
that it stops far short of the nostrils. Sometimes, in addition to this 
bare spot, two other minute spots occur, one on each side of the 
former but closer to the nostrils. The lips of elk are covered with hair 
right up to the edge of the mouth. 

On the legs, especially in their lower half, the hair is short and 
not wavy, slightly arcuate, resilient, and very strong, especially on 
the anterior side of the legs. This represents an adaptation in an 
animal which for most of the year moves through deep snow. 

The coloration of the trunk, the upper portions of the legs, the 
neck, and much of the head of an adult animal is a uniform, dense, 
brownish-black or nearly black. The tip of the muzzle is light gray, 
even whitish. The coloration of much of the limbs, roughly from the 
middle of the shin and foreshoulder, is light gray, sometimes nearly 
white with a silvery shade, and contrasts sharply with the overall 
color of the uunk." Speculum absent. 

There are no significant differences in the color of the summer 
and winter coats of elk. Only a single molt occurs, in spring, but it 
extends over most of the summer. Commencing in April molt is 
most intense in May and June and remains of the worn-out winter 
coat are still evident even in July. The complete short summer coat 
(August) is extremely dark, almost black, and shiny. Later, due to 
hair growth, the color gradually lightens by winter, becoming more 
brownish and dull. The wear and tear of dark hair tips in winter also 
contributes to the appearance of brownish shades. There are no 
sex-related color differences. 

The coloration of newborn elk during their first few months (up 
to the first winter pelage) is a uniform dark or rusty-brown all over 
the body. The anterior portion of the muzzle and the legs are also of 
the same color. Sometimes a dark band occurs along neck and in the 



206 ^Yie fur from the legs of reindeer, maral, and elk — the so-called "kamus" — is 
used to line the lower surface of taiga skis. This facilitates their movement and 
prevents them from slipping backward ("kamus" are fixed with the hair directed 
backward). Elk "kamus" is particularly strong. 

^"'in winter frosts, especially in hazy light, the light-colored legs of the animal on 
snow are very poorly visible and during slow movement the impression of a dark- 
colored animal floating in air is common. 



314 

shoulder region on the back. Spotted coloration is never seen in the 
young. 

In adult bulls body length reaches 300 cm, height at withers 225 
to 235 cm, length of ears about 26 cm, and tail 12 to 13 cm. Weight 
reaches 570 kg. ^°^ Antler span up to 150 cm and weight up to 20 kg; 
even larger antlers have been recorded, however. 

Overall length of skull up to 620 mm and zygoma width up to 
240 mm.^°^ Cows invariably smaller and more slender. 
226 Body length of newborn about 90 cm and height at shoulders 70 
to 90 cm. Weight 13 to 16 kg. In the first autumn of life the young 
weigh about 100 kg, which rises to about 200 kg by the end of the 
first year. (V.H.) 

Taxonomy 

Genus Alces contains only one species. 

Geographic Distribution 

Alces are distributed in the forest zone, partly in forest-steppes, and 
the edge of steppe zones in North America, Eurasia, and the 
Caucasus. 

Geographic Range in the Soviet Union 

The range in the Soviet Union represents the major part of the range 
of the species and an overwhelming portion of the range in the Old 
World. It covers almost all Soviet territory except semidesert and 
desert regions and the mountains of Middle Asia. 

The range of elk is of special interest in some respects. Its major 
changes over the historic period and the particularly extensive 
colonization of the animal in the last decade are of utmost theoreti- 



These data are from Flerov (1952). According to others (Buturlin, 1934; Zhitkov, 
1914; Kulagin, 1932) the maximum size of Russian elk is somewhat greater — body 
length may exceed 300 cm, height at shoulders 240 cm, and weight 620 or even 655 kg 
(40 poods). These maxima, assuming them to be fairly accurate, represent rare excep- 
tions and pertain mainly to animals of northeastern Siberia, which are particularly 
large. Some data were collected at the end of the last century when elk at many places 
were of far greater dimensions than at present. 

"*" See characteristics of the genus for skull features. 

"'"' Morphological and morphomeuic data from Kulagin, 1932; Buturlin, 1913 and 
1934; Zhitkov, 1914; Yurgenson, Kaplanov and Knize, 1935; Flerov, 1952; and other 
sources, including original material. 



315 



cal importance from the viewpoint of zoogeography and ecology. 
Contraction and expansion of a range, movements and pulsations of 
its boundaries, forms of colonization, occupation of new territories, 
and several other aspects of general zoogeography are vividly 
revealed in the distributional history of elk. 

From an ecological point of view a study of the distribution of 
elk and the dynamics of their range provide extremely interesting 
possibilities for delving into the affinity of the animal to a given 
topographic region, its specialization and ecological adaptability, 
range expansion into new areas, and several other features, for 
example, the origin of regular seasonal migrations in an otherwise 
resident animal. Of special importance is the coexistence of a large 
ungulate with man in arranging the creation and development of 
cultivated landscapes. 

A study of the range of elk is also of practical interest. Elk is an 
important species of commerce and represents one of the most prom- 
inent objects of hunting in the world fauna. The prospects for utiliz- 
ing it for these purposes are extremely good. The fate of elk and 
changes in its range are of exceptional interest from the point of 
view of conservation. It can be confidently stated that there is no 
other large animal whose conservation despite all handicaps has 
provided such extensive and amazing results. Even though many 
other species facing extinction have been successfully conserved and 
their population and range enlarged, achievements have nowhere 
attained those in the case of elk. The results of elk conservation are 
only comparable to those achieved with saiga antelope (see below). 
From experiments conducted outside the USSR the effect of conser- 
vation of elk can be compared to some extent only to that achieved 
with the Pribilov fur seal herds. However, in the latter all the prereq- 
uisites of success were relatively simple. The same can be said for the 
conservation of Russian sable. 

The view that elk represent typical inhabitants of dense forests, 
with a preference for swampy taigas, and to a lesser extent mixed 
taigas in dense forests, prevailed for many years. Similar views were 
expressed about its range. In the last decade historical and archaeo- 
logical finds have shown that elk lived far from true taiga. They 
228 were extensively distributed not only in the zone of mixed and decid- 
uous forests, but in the forestlands of forest-steppes and in bottom- 
land deciduous forests, and even extensively dispersed in the steppe 
zone. It has also been established that elk inhabited the montane 
forests of the Caucasus comparatively recently. 



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317 

Nearly 40 years of legal conservation have further revealed that 
elk, long regarded as a species facing almost total extinction and shy 
of man ("fugitive from civilization") are, in fact, animals of high 
vitality capable of extensive distribution. In the absence of direct 
persecution elk are capable of living, and do so, in the immediate 
proximity of man in densely populated areas and the environs of 
large cities. Elk can even be designated somewhat "anthropophi- 
lous". Nevertheless they are very readily susceptible to extinction. 

Right now, with the population and its density having risen 
greatly and standing high almost everywhere, the range of elk has 
notably enlarged. As in the distant past, the animal once again 
inhabits forest-steppes, has penetrated even the steppe zone, entered 
the forest- tundra, and even the tundra in the north. This confirms 
well a characteristic of elk, established on the basis of a study of its 
former distribution, that ecologically it is an extremely adaptable 
eurytopic species and not a strictly stenotopic inhabitant of taiga. 
The confinement of elk to dense taiga may be regarded as mainly 
due to human persecution. While this zone can be considered an 
ecological refuge of the species, it is nevertheless quite clear that it 
does not represent an optimum environment for them. 

As far as the range of elk and its dynamics are concerned, dense, 
tall forests are less favorable than sparse forests with meadows, 
swamps, and burned-over and felled forest sections. Such areas ensure 
important winter food-shrubs and twigs. Burned-over and felled 
forest sections, overgrown with deciduous species, mainly aspen, are 
particularly favorable in this respect. 

Parallel to conservation, such habitats are one of the important 
factors determining the expansion of range and growth of elk popu- 
lation. Extensive and often continuous felling in the last decade over 
much of the range of elk radically altered the forest landscape into a 
zone favorable for its habitation. Herein lies one of the vital factors 
responsible not only for the rise of population but also for the 
"anthropophilous" character of elk. 

In this connection it is very significant that in western Siberia the 



^" Since the 1950's elk have been sighted often in the environs of Moscow and 
especially Leningrad, where they are sometimes held in, and live around, parks. 



121 Fig. 70. Geographic range of elk (Alces alces L.) in the USSR in mid-1950's 

(scale in km). 

1 — boundaries of region of permanent habitation; 2 — boundaries of regions of visita- 
tions. For details see subsequent maps. V.G. Heptner. 



318 



10 16 20 26 30 36 40 46 60 66 60 66 70 76 




200 О 200 400 600 km 
' ■ ■ ■" ■ ■ ' — 



-9—?- 2 — p^—r 5 
Г-» 3 



319 

population density of elk reaches the highest level in the southern 
zone where taiga is sparse and acquires some forest-steppe features 
(LP. Laptev, 1958). It has been show^n (Sludskii, 1953; and others) 
that, in the past (eighteenth century and before) elk were abundant 
in the forest-steppe and a common animal. Thus the widely preva- 
lent view that elk is a northern species cannot be justified. 

A characteristic feature of elk, judged from some data collected 
over the last two-and-a-half centuries, is that its range changed 
extensively over a comparatively short period (contraction and 
expansion) and the boundaries shifted over long distances. In some 
cases and for certain parts of the range the reasons for these rapid 
fluctuations — usually man-induced — are quite clear, while else- 
where the causes of the phenomenon are not known. On the whole 
this phenomenon has not been thoroughly studied and is under- 
stood only for part of the range (European part of the USSR and 
western Siberia). However, there is reason to believe that this pheno- 
menon was quite widespread not only in a territorial sense, but also 
in time. 

Different sections of the range over the historic period are de- 
scribed below. 

229 Geographic Range in the European Part of the USSR 

Reconstructed range and range in seventeenth and eighteenth cen- 
turies. The range in the period of its maximum distribution (Fig. 71) 
covered almost the whole European part of the USSR, the forest- 
steppes, and much, if not all, of the steppe zone. 

On the basis of the few historical records available the southern 
boundary of the range of elk was probably as follows. Commencing 
in the west at the boundary of Russia southwest of Kishinev, from 
slightly south of this town, it crossed the Dnestr and went to the Bug 

230 Fig. 71. Some changes in boundaries of distribution of elk {Alces alces L.) in 
the European part of the USSR over the historic period and up to the 
twentieth century (scale in km). 

1 — reconstructed southern boundary of region of fairly permanent habitation over 
historic period. Individual transgressions evidently extended beyond this zone. Bound- 
aries in the Caucasus simplified (for more details see Fig. 73). Arrow indicates 
■ probable migration from Volga into Kuma; 2 — probable position of southern bound- 
ary in seventeenth and eighteenth centuries; 3 — southern boundary around 1850, the 
period of utmost recession before mass colonization of elk in the south in latter half of 
century. Some individual pockets of habitation falling south of the northern bound- 
ary shown (Koppen, 1883); 4 — probable northern boundary at time of its maximum 
recession to the south in the middle of the nineteenth century (around 1850); 5 — 
southern boundary of advancement of range southward during latter half of the 
nineteenth century (early 1880's; Koppen, 1883). V.G. Heptner. 



320 

somewhere in the region of Voznesensk. From there it proceeded to 
the Dnieper in the region of Dnepropetrov (probably even more 
south) and proceeded toward the Northern Donets, probably in the 
region of Izyum. Later, it evidently proceeded along the Donets to 
Don and descended along the latter to the Sea of Azov. The range 
also covered the Don valley above the mouth of the Northern 
Donets. Evidently in the region very close to the Don and Volga the 
boundary bypassed on the north the semideserts on the right bank of 
the Volga and crossed to the Volga; the range descended thus in the 
form of a narrow extension along the Volga valley down to the 
Caspian Sea. 

The range (zone of permanent habitation) evidently did not 
cover the zone of arid steppes and semideserts between the Volga and 
Ural [rivers]. From the Volga valley the southern boundary crossed 
to the Ural valley, probably roughly along the line from Stalingrad 
to Ural'sk, following the northern boundary of the semidesert. The 
range descended south along the Ural valley in the form of a projec- 
tion. The maximum distribution of the species is not known but the 
possibility is not ruled out that it extended to the sea. 

There is no doubt that in the forest-steppes of long ago, with 
innumerable large forest massifs and multiple small ones, elk were 
distributed widely and fairly uniformly. In the steppe region, how- 
ever, the range was not continuous. Elk were confined to some indi- 
vidual forest massifs and to bottomland deciduous (gallery) forests 
along rivers which cut into the steppes. There is no doubt that elk 
were thus not distributed exclusively along the Don and Volga. We 
can assume they were widely distributed south of the line shown 
[map] between the Danube and Don. They evidendy reached the sea 
through valleys of not only large rivers such as the Dnestr, Bug, and 
Dnieper,^'^ but also along much smaller rivers such as Molochnaya, 
Mius, and others, on which also there were bottomland deciduous 
forests in the distant past. Indubitably they emerged into open 
steppes for grazing while moving from one island to another, and 
during migrations were generally confined to open steppes which, 
in olden times, were far richer in tree and shrub food than in the 
Recent period. A similar picture is currendy seen in Kazakhstan and 
the Trans-Volga where the animals not only perform major migra- 

^'^ According to Kirikov (1955) and unpublished data furnished by S.V. Kirikov 
and N.K. Vereshchagin. 

^'^ Elk have been reported in the lower courses of the Dnestr (Kirikov, 1955; map) 
at Ol'vinsk on the shore of the Bug estuary (1000 A.D.) and on the shore of Kuyalnits 
estuary (Il'insk, 1000 B.C.; Pidoplichko and Topachevskii, 1953). 



321 

tions (up to 100 km) across open steppes but also enter semideserts. 
There is every reason therefore to assume that the range of elk 
between the Danube and Don extended up to the shores of the Black 
and Azov Seas. It is highly possible that even in Trans-Volga its 
boundary lay more southward and Uzen' also fell within the range. 

The situation described above prevailed in the historic past, 
probably in the early part of the last millenium and at places later, 
probably up to the sixteenth or even seventeenth century. Informa- 
tion is available about the occurrence of elk in the lower course of 
the Don from 800 to 1000 A.D. (ancient township of Tsimlyansk). It 
is possible that the range was just as extensive, or almost so, even at a 
much later period. Pallas has written (1811) that around 1800 elk 
thrived throughout Russia from the White Sea to the Caucasus, i.e., 
evidently lived even in forest-steppes and steppes. 
231 There is no information whatsoever about the northern bound- 
ary of the natural range over the historic period. Undoubtedly, 
however, it could not have been farther removed than the boundary 
at the middle of the present century, i.e., the southern boundary of 
the tundra and the sea coast (see below). This supposition is none- 
theless tentative. 

In the seventeenth and eighteenth centuries the boundary evi- 
dently passed along the northern part of the steppe zone or along the 
southern rim of the forest-steppe. From Moldavia to the Northern 
Donets it deflected slightly north or generally corresponded to the 
previously mentioned Kishinev-Voznesensk on Bug-Dnepropetrovsk- 
Izyum on Northern Donets line. Elk were present even in the Black 
Forest at the watershed between the sources of Ingul and Ingul'ts 
(about 1740), and below Izyum (about 1667). However, to the east the 
range boundary rose considerably northward, crossed the Don at 
about 50° (Boguchar region), and ran along the Khoper in the 
Novokhopersk region or slightly more southward (elk were present in 
Tellermanov Grove around 1657). From there the range boundary 
proceeded toward the Volga, probably straight eastward. It is highly 
possible that in the forests of the Khoper and Medveditsa elk were 
present even more southward. No information is available on the 
Trans-Volga region. 

In the Caucasus (Fig. 72) the region of elk occurrence adjoined 
the Russian plains. A connection occurred through the lower 



"'"According to data of S.V. Kirikov. 

^'^The first reference to the presence of this species in the Caucasus is found in 
Pallas' works (1811). He states that elk were present "omnique Rossiam, aMari Albo 



322 




232 Fig. 72. Former distribution of elk {Alces alces caucasicus Ver.) in the Caucasus (scale in 

km). 

1 —probable range in historic period; 2 — references in literature for the early nineteenth 
century; 3 — dzvars in which elk were found; 4 — habitations on Urup River; 5 — kitchen 
middens of eighth to thirteenth centuries; 6 — remains of fourth to first century B.C.; 
7 — Upper Palaeolithic remains. Arrows indicate probable courses of colonization and 
migration of elk in the Caucasus (original by N.K. Vereshchagin, simplified and with 
supplementary information from S.V. Kirikov). V.G. Hepmer. 



course of the Don and the Azov region of the northern Caucasian 
steppes. More to the east, where dry steppes and semideserts he, no 
permanent Unkage of the range is evident. However, it is wholly 
possible that the animals migrated along the Caspian Sea (from the 
Volga estuary) but due to the recession of the southern boundary of 

usque ad Caucasum". Considerable controversy ensued but by and large Pallas was 
deemed wrong (see Koppen, 1883; Dinnik, 1914; Satunin, 1910; and others; more 
recently, Kulagin, 1932). The former occurrence of elk in the Caucasus was conclu- 
sively proved however by N.K. Vereshchagin and O.I. Semenov-Tyan-Shanskii, 
1947. They found elk skulls in Ossetian dzvars [ritual sites] at Lezgor and Digorized in 
Urukha River basin (Vereshchagin and Semenov-Tyan-Shanskii, 1948; \'ereshchagin, 
1948; Vereshchagin and Naniev, 1949; Vereshchagin, 1949). In the light of these data 
not only the assertion of Pallas but also data from other early authors, such as biologists 
(Gmelin, Georgi, Clark, Middendorff), as well as hunters (Polferov, 1894), and 
references in Ossetian folklore have acquired special significance. 



323 

the range in Volga-Don region, described above, this continuity was 
interrupted. 

Outhnes of the Caucasian elk range can be reconstructed only in 
an extremely tentative form, largely hypothetical, and only for the 
last century. Elk evidently lived in the steppe parts adjoining the Sea 
of Azov where reed thickets were abundant and bottomland decidu- 
ous forests were present not only in the Kuban delta but also along 
streams flowing into the sea. Later the animals moved into the 
whole of the forested western extremity of the mountain range and 
the forests on its northern slope and foot-hills. In the past these 
forests penetrated far into the plains and were closely associated with 
forests lying on the Stavropol highlands. In the east elk probably 
lived on the plains, in forests of the Terek basin, and in the lower 
reaches of the rivers their range probably reached, or almost reached, 
the Caspian Sea. 

In the Trans-Caucasus the range of elk extended in the form of a 
belt along the Black Sea coast and slightly enlarged in the basin of 
the Rion and other rivers of this part of the country without, how- 
ever, reaching the Kura basin. The possibility is not excluded of the 
range crossing slightly into Turkey in the region of the lower 
reaches of the Chorokh or still more southward. 

The range described above is that of the fifteenth to the early 
nineteenth centuries. Elk were extensively destroyed in the Caucasus 
in the seventeenth and eighteenth centuries and consequently the 
range shrank notably. The last elk, judging from skulls in the sanc- 
tuaries of the central Caucasus, were probably killed in the early 
nineteenth century. In the lower reaches of the Kuban, between the 
main channel and the tributary at Kalaus village, elk were sighted 
even in 1800 (Clark 1810; cited by S.V. Kirikov) and evidently were 
not exceptionally rare there. 

The period of penetiation of elk from the north into the Cauca- 
sus should perhaps be placed at the very end of the Pleistocene. In 
the Upper Palaeolithic period elk were evidently dispersed within 
the region described. In the lower Don at Tsimlyansk elk remains 
were found in the ancient township dated 800 to 1000 A.D. (they 
were absent in Sarkela from 900 to 1100 A.D.). It is difficult to state 
precisely when the contact between the Caucasian and northern 
range was broken. Apparently it occurred at the commencement or 



^'*Two young elk at Kalaus were extremely tame; Clark (1810) was informed that 
many wild elk could be seen in the steppes in spring. 

^'^The elk range in the Caucasus is based on articles cited before and on the 
unpublished data and map of N.K. Vereshchagin. 



324 

in the first half of the last millenium. This is borne out by the 
taxonomic isolation of Caucasian elk, which represent a different 
subspecies. Furthermore, there has never been a full merger between 
Caucasian and Russian populations. This also may have influenced 
differences in the morphology of the Caucasian form. 

Range in nineteenth and twentieth centuries. In the first half of 
the nineteenth century a very sharp, almost catastrophic reduction 
in the range of elk occurred and its southern boundary receded far 
north. Compared with the boundary line shown for the 1700's, by 
the mid-1800's the boundary had shifted at places 450, 600, or even 
1 ,000 km. Recession was particularly far in the central regions and 
less in the west and east. 
233 By 1850 the southern boundary of the elk range in the European 
part of the USSR formed an irregular line running roughly through 
the following places: mouth of the Neman, upper course of the 
Pripyat, middle course of the Sluch, mouth of the Pripyat, upper 
course of the Desna, Kalininsk, Rybinsk, Bui (northernmost point), 
lower course of the Unzha, middle course of the Vetluga, the Volga 
east of Vetluga mouth, region of the mouth of the Vyatka, mouth of 
the Beloe, middle course of the Ufa, sources of the Ufa and Ural 
[river], and eastern slope of the Urals [mountains] at roughly 56° N 
lat. Outside this boundary (south of it) there were two small islands 
of habitation of elk, east of Moscow in the forests of Vladimir district 
and between the lower courses of the Sura and Volga north of 55°. 
Other pockets in which elk survived could also have existed 
(Koppen, 1883; boundary description as depicted in Koppen's map). 

In the nineteenth century a notable southward recession of the 
northern boundary occurred, which chronologically coincided with 
a similar phenomenon along the southern boundary, already de- 
scribed, for the first half and middle of that century. The northern 
boundary in the west (Karelia) at that time ran roughly between 64 
and 65° N lat. and in the east proceeded from the source of the 
Vychegda, passed to the south of the Pechora basin and descended 
southeast almost to 60°, intersected the Urals there, rose slighdy 
northward passed to the north of the Sosva, and ran into the upper 
Lozva (Nasimovich, 1955). Distribution in the area between White 
Sea and the source of Vychegda is not known. This depression of the 
northern boundary of the range was associated with the correspond- 
ing phenomenon in Siberia (see below). 

Thus by the middle of the century the range of elk had shrunk 
sharply because of the extremely pronounced shrinkage of the south- 
ern boundary and to a lesser extent of the northern boundary as well, 
which compared to the period of maximum enlargement lay at places 



325 

even 500 to 600 km more southward. 

The reasons for this reduction in range are not clearly under- 
stood because they were inadequately studied. The human factor 
(hunting), of course, played a significant role. According to some 
authors (Nasimovich, 1955) it was the decisive factor.^'^ However, it 
does not seem credible that this factor alone could have caused the 
extensive shift of boundaries which occurred fairly synchronously 
over an immense expanse from the Baltic region to the Urals, and 
farther to the east, a shift which compelled the animal to move out 
of an extensive territory extremely favorable to its survival. The 
chronology of this phenomenon has also not been traced com- 
pletely. It is difficult to state when it commenced but without doubt 
it developed over some decades and the boundary described formed 
as a result of steady change which was apparently quite rapid. In 
any case, at several places where elk were formerly quite common 
the local populace and even hunters had forgotten this animal by 
the middle of the century. When later a southward advancement of 
elk began anew, quite often, for example in some parts of Simbir 
province (Ulyanov district), it was a totally unknown animal to the 
local people. In the historical perspective of the species (even over 
the last few centuries), the range shrinkage described above should 
be considered rather rapid. This was a spontaneous "compression" 
of the range with very few parallels. 
234 After 1850, when the contraction of the range had evidently 
reached maximum, reenlargement commenced in the south. Elk 
began reappearing in places where they had been absent for several 
years. Animals from northern forests often settled down at fairly 
long distances from their original habitats. They colonized new pla- 
ces and multiplied, often quite rapidly. Population increments were 
due not only to reproduction, however, but also the arrival of new 

The main reason for the intense persecution of elk in the first half of the last 
century was not the animal's meat. Under sparse population conditions it was of no 
commercial importance. Hunting of the animal was stimulated by the use of elk hide 
by the Government which, commencing from the time of Peter the Great, used them in 
uniforms for some regiments and elk chamois in riding breeches and jackets. The use of 
elk hide finally ceased in the nineteenth century because of the sharp reduction in 
population and range of elk. At the commencement of the twentieth century only one 
guard regiment was outfitted in a uniform made from elk hides and that, too, for 
ceremonial purposes only. 

Incidents of occurrence of European bison in B. Nizhegorod province reported in 
literature in the middle of the last century evidently relate to captures of elk. At that 
time elk were so rare in the province that the local people did not recognize them and, 
having forgotten the name "elk," called these animals "builo" [a type of bison] 
(Koppen, 1883). 



326 

groups of animals in considerable numbers. 

The southern boundary of the range advanced eastward and west- 
ward over relatively small distances, up to 100, 150, and 200 km, but 
in the central regions up to 500 to 600 km. By 1880 it ran roughly 
along the following places: upper reaches of the Sluchi and Dnieper 
slightly north of Kiev, Chernigov, Desna, slightly south of Bryansk, 
slightly north of Orel, upper reaches of the Voronezh, slightly north 
of Tambov, slightly south of Penza, slightly south of Ulyanov, and 
lower reaches of the Beloe (Koppen, 1883; the above line is based on 
Koppen's map). Southward advancement of elk continued even 
later, in the 1880's and the 1890's to a lesser extent. At that time the 
population of the animal in colonized regions also increased. 
Finally, by the early twentieth century elk had become quite com- 
mon at many places and were even abundant in some (B. Simbir 
province). Data concerning these phenomena are quite sketchy, 
however. 

The southward movement was rapid, more rapid and more per- 
ceptible anyway than the boundary recession. At some places coloni- 
zation was evident and rapid because of migrations over long 
distances and the unexpected appearance of the animals; fairly large 
numbers appeared suddenly in some areas (Alatyr basin, Simbir 
province, now Ulyanov district). 

The reasons for the foregoing phenomena are not clear. It is 
possible that an intense felling of forests in the central provinces, 
commencing after the agitation of local landholders in the 1860's 
(abolition of land rights in 1861) played an important role. These 
felled clearances led to the formation of large areas with young 
saplings, i.e., to the formation of an environment extremely favora- 
ble for elk. This, however, could not have been of decisive impor- 
tance and at best would only have influenced its course in some 
individual limited sections. The environment in the southern forest 
zone had always been favorable for elk. Moreover, at many places a 
significant number of elk appeared even in the absence of such 
clearances under conditions which did not at all vary from the origi- 
nal ones (forest along the Sura River). Evidently the settlement of 
elk in the south was stimulated mainly by conditions prevailing in 
regions from where migrations took place. There is no doubt that 
the colonization of elk in the south must have been preceded by a 
significant increase in their population in the north. It is possible 
that forest fires compelled elk to move out from some regions. 

^^"These and other data cited above for Simbir province were communicated to the 
author by S.A. Buturlin some time ago. 



327 

Range at commencement of twentieth century and in the 
1920's. information is inadequate for precisely demarcating the 
boundaries of the elk range in the early twentieth century. By then 
the southern boundary corresponded fairly well to that described for 
1880. In the upper reaches of the Don and Oka, and probably also at 
other places, it possibly lay somewhat more northward; in other 
areas it may have occupied a more southern position (Trans-Volga, 
Ural, and probably in the west). In any case, early in the century elk 
were widely dispersed in the forest zone and reached quite far into 
the south. Their population was fairly high, for example in the 
provinces of Vladimir and much of Ryazan' and Moscow as well. 
There the animals lived in forests directly adjoining Moscow, which 
are now markedly reduced (Pogonno-Losin Island, Novogireevo, 
and Losin Island; V.G. Heptner). This was the situation until the 
mid-1910's. 

235 At the end of World War I and in the years of the Civil War the 
elk population was drastically decimated by unrestricted poaching. 
In much of their range in the European part of the USSR elk became 
rare and faced total extinction. Banning of elk hunting, decreed in 
the very first year of the Revolution, exerted some influence only by 
the mid-1920's. The southern boundary of the range again receded 
greatly northward. The southern boundary of the range in this 
period of maximum depletion has not been established. It is fairly 
well known for 1928, when some signs of restoration of the elk 
population were observed or, more correctly, population reduction 
and decrease in the range were checked. It is even possible that at 
some places the range enlarged somewhat, of course very impercep- 
tibly, and the 1928 line can generally be considered close to the 
boundary of utmost depletion of the range in the 1920's. 

This boundary (Yurgenson, 1935; Danilov, 1951) commenced in 
the west at the former Russian state boundary, slightly east of the 
lower reaches of the Sluchi, and ran east slightly north of 51° N lat. 
Short of 30° E long, it turned steeply north, later west, passing to the 
west of the Minsk district, and again turning east crossed the upper 

236 reaches of the Berezinsk. From there it again turned north, left much 
of the upper reaches of the Western Dvina out of the range, and 
running southeast crossed the very source of the Dnieper. Farther 
away, forming a narrow loop southeast of 35° E long, (toward the 
Volga) it turned sharply southwest and, describing an arc, crossed 
the Desna twice — initially at its source (southwest) and later in 
Bryansk region or slightly south of it (on the east). 

From this place the boundary line turned initially northeast and 
crossed the Oka slightly southeast of Kaluga; Jater it proceeded 



328 

slightly north of the Oka and left the Oka roughly at 55° N lat. 
(slightly north of the mouth of the Moksha). From there the range 
formed a long and narrow extension along the Tsna southward 
roughly up to Tambov. The eastern boundary of this extension ran 
through the midcourse of the Moksha in the vicinity of the mouth of 
the Oka. Between the mouths of the Oka and Sura the boundary ran 
slightly short of the left bank of the Volga. From there the range 
formed a large triangular projection to the south. Its western bound- 
ary ran initially along the Sura and later left toward the interfluve 
region of the Volga and Sura, slightly north of the source of the 
Sura; the eastern boundary from there turned toward the Volga 
north of Ulyanov. Later, the boundary proceeded initially southeast 
and then north toward the mouth of the Beloe to Kam. Continuing 
farther, initially somewhat northward along the left bank of the 
Kam, it later turned east steeply and, crossing the upper reaches of 
the Ufa, ran northeast into Siberia through the source of the 
Chusovoi. 

The northern boundary of the range in these years, as in the 
much earlier period, is very poorly known and can only be drawn 
tentatively. In the Kola peninsula elk were evidently not distributed 
in the north beyond the forest boundary and did not enter the forest- 
tundra toward the Barents Sea.^^' Toward the White Sea in the east 
the boundary probably extended up to the latitude of the arctic 
circle. Thus elk were encountered in the southern half of the penin- 
sula. On the mainland the northern boundary apparently extended 
along the shore of the White Sea and probably reached, or almost 
reached, the lower reaches of the Mezen'. The range encompassed 
the entire basin of the middle and upper reaches of that river. In the 
Pechora basin the range boundary descended sharply southward and 
evidently went around it to the west and crossed somewhere in the 
upper reaches, probably around 62° N lat., running toward the 
Urals at 61° to 62° N lat.^^^ 

Around 1930 the boundary ran roughly along the Ponoi, crossed mid-course of 
Vorona, and ran across the Pulozero at Notozero (Semenov-Tyan-Shanskii, 1948). 

^"^The boundary of the elk range in the Pechora basin and some parts of the Urals 
underwent considerable changes in the last century. Sometimes it moved northward 
and at other places significantly southward, until elk appeard on the Pechora only as 
transgressors. The animal population also fluctuated sharply. 

In the Pechora basin the boundary fluctuation in the last century was evidently 
associated very little with the extinction of the animal and depended more on the 
condition of food sources (availability of young saplings of deciduous species). 
Animals moved from this region and colonized elsewhere. Because of scanty 
information it is not possible to draw a satisfactory chart of the phenomenon (some 
data are provided by Teplov and Teplov, 1947; and Nasimovich, 1955). 



329 

Geographic range in 1930's and 1940's and present bound- 
aries. Following the range contraction described above, a gradual, 
initially very weak and later increasingly rapid expansion, occurred 
from the end of the 1920's and at some places probably somewhat 
earlier. It proceeded south and partly north and northeast (Pechora 
basin and the Urals). This phenomenon is the result of planned 
conservation and attendant increase in population. In the European 
part of the USSR from 1920 through 1950 elk increased roughly 15 
to 20 times (Danilov, 1951; see "Biology" below). 

The southward expansion of the range in the European part of 
the USSR proceeded at the rate of 200 or even 400 km over 18 to 20 
years. On the average the boundary there in those years advanced by 
120 km and the range enlarged southward by over 400,000 km.^ Thus 
in the 1930's and 1940's the range in the forest zone was reestab- 
lished, completely colonized, and animal density quite high. From 
the 1940's colonization of forest-steppe zones commenced and pene- 
237 tration into steppe zones was observed at places; some animals 
moved even into the semidesert zone. 

By 1947 the southern boundary of the range had undergone the 
following more important changes. The area of the above-described 
northward curve of the boundary into the basin of the upper 
Dnieper and upper reaches of the Western Dvina was greatly reduced 
in the west. Its northernmost point lay roughly in the Mogilev 
region. In the region between the Desna and Tsna the range on the 

The occupation of new places by elk was initially in the form of occasional 
visitations by isolated animals which later increased both in number of animals and 
frequency of visitations. The animals gradually settled in the new place and began 
multiplying. In the forest zone this process was hardly perceptible; transgressions were 
usually over small distances and advancement of the range southward very gradual. 
During the occupation of southern zones — forest-steppes and steppes — occasional tran- 
sgressions were also observed, usually in "spurts'" over long distances (for example, 
tens and hundreds of kilometers in the open steppes). The animals often happened 
into a totally inhospitable environment (sometimes in a town) and usually died. Gradual- 
ly these visitations became increasingly frequent and some animals, finding fairly 
suitable conditions (islands of forest in the steppes), settled down. Sometimes col- 
onization occurred in alien zones — steppes, semidesert, tundra, along bottomland 
deciduous forests and groves, and even shrub thickets. 

At some places regular seasonal (summer) migrations of elk into tundra occurred 
and the animals were sometimes trapped into wintering at places north of the region 
of permanent winter residence. As a result of these factors it is difficult to distinguish 
the region of permanent habitation from the region of transgressions in several cases 
over large territories, all the more so because the process of occupation of new areas 
is still continuing. Expansion of the range of elk and the mode of expansion are of 
considerable theoretical interest for understanding the phenomenon of colonization 
in general. 



330 



20 26 



ЭО 36 40 46 



66 во 




238 Fig. 73. Southern boundary of distribution of elk (/4 Zr«aZceiL.) in the European part 
of the USSR in 1958 (scale in km). 

1 — southern boundary of region of fairly permanent habitation; 2 — southern 
boundary of region of visitations from 1955 to 1958 (in Trans-Volga some 
transgressions which took place in the very early years have also been taken into 
consideration). Circles on line represent some important definite points referred to in 
the text; 3 — points of particularly distant visitations (1 — Gorodok region; 2^ 
Solobkovet region; 3 — Novopokrov region; 4 — Sambek). Question mark on Ural River 
indicates that elk probably spread farther south; 4 — southern, western, and part of 
northern boundary of forest-steppe region; 5 — southern boundary of steppe zone in the 
Crimea and the Caucasus; б — northern and western boundaries of semidesert and 
desert zone. Boundaries of natural zones shown schematically from data of S.V. 
Kirikov. V.G. Heptner. 



whole advanced somewhat southward, and that too very insignifi- 
cantly. But then, between the Tsna and Volga and along the Volga 
the boundary shifted southward very extensively, although not 
entirely uniformly. The curvature of the boundary in the north 
between the Tsna and Sura almost disappeared and the boundary 
descended along the Volga almost to Saratov. The entire boundary 
advanced particularly extensively in the south into the Trans-Volga 



331 

and Urals. By 1947 it had already extended from Samarsk Luka to 
the southern Urals, where it stopped only slightly short of the Ural 
River valley in its meridional course. 

The southward advancement of the boundary, compared with 
the 1928 position, was 100, 200, or even 500 km at some places. At 
other places, contrarily, it did not advance at all. This was particu- 
larly so at places where the range had reached the natural topogra- 
phic boundary even before, but primarily where further advancement 
was hampered by human intervention (some details of the boundary 
are shown in Fig. 74). 

Adequate information is not available about the northern bound- 
ary of the range of elk in the period under description (1928 to 
1947). It is known, however, that the population of the animal 
increased even in the north and the range enlarged in that direction 
also. In the northern Urals, by that time, it evidently reached 65° N 
lat. or approached it. Probably the advancement in the Pechora 
basin was also roughly of the same magnitude or slightly less. 

After 1947 the elk population and its density in the European 
part of the USSR continued to increase and the southern boundary 
of the range continued its progressive advancement southward. The 
animals occupied an increasingly greater area of the forest-steppe 
zone. By 1952 elk in the Urals descended so far southward that they 
colonized not only the entire mountainous district, but also the Ural 
[River] valley from Orsk to Orenburg (55° E long.) and extended 
southward even beyond the river. 

In 1952 the southern boundary of fairly permanent habitation of 
elk in the European part of the USSR traversed from Orenburg 
through Buzuluk Bor to Kuibyshev to the Volga and farther down 
along the river roughly to Saratov. Along the left bank of the Volga 
in this section elk were encountered on the Chapaevka River 
(Mocha) above Chapaevska, i.e., south of Kuibyshev and in the val- 
ley of the Great Irgiz opposite Vol'sk (west of Gmelinovka station; 
Martynov, 1951; Stroganova, 1951). From Saratov the boundary ran 
southwest to Shirokii-Karamysha on the Medveditsa at the southern 
boundary of the Saratov district and from, there up along the river 
toward Petrovska and then to the west toward Rtishcheva and 
239 Makarova. It then turned upward along the Khoper to the Penzena 
district and later along the Tsna basin in the south almost to Tam- 
bov. From there the boundary proceeded toward Ryazan', through 
the source of the Don at Orel, then to the southwest toward the 

^^''The boundary of distribution of elk in 1947 is based on data from D.N. Danilov, 
1951. 





10 

1 ji 


15 20 26 30 
■ - 1 —ЛИ ж I 1= 


36 


40 46 50 55 во в6 70 76 


" 


6 




»^~"* 






65 
60 




3 


к ^ 


6 




W bS. R^ I 




5 

5C 








55 

50 




^ 


'"irr^-^d-Zi 


s 












^ 






'i+ 


jf 1^=?"^ L-.^ 


5^ 


^'^vl^ 


f\ (^ 


48 






М 




V^. vi^i-W: e \ ^Л-г~ 




1 


j?^^^fe 




к t 








J/ ^-^f— -^ 


Ч 


л^_гг^ 




(J 




г 


la^.^^^ 


40 




^■^^ / t;^ 


^^ 


^'vj y^A - 




200 


200 400 600 km 












Ir-r-r- 4 

-.— 2*5 
3*6 








^^ 












40 


46 





238 Fig. 74. Boundary of geographic range of elk (Alces alces L.) in the European part of 
the USSR and changes in it in the first half of the twentieth century (up to 1952) (scale 

in km). 

1— southern boundary in 1947 (from Danilov, 1951); 2— same in 1928 (from Danilov, 

1951); 3— same in 1952 (V.G. Heptner); 4— northern boundary in 1920's; 5— some 

particularly deep transgressions in the 1930's, 1940's, and beginning of 1950's (1951 and 

1952); 6 — transgression toward Kara Bay. V.G. Heptner. 



333 

midcourse of the Desna and in the north to Roslavl' and Mogilev. 
From there it turned south to the mouth of the Sozh on the Dnieper 
and the lower reaches of the Pripyat. Having encompassed a signifi- 
cant part of the southern tributaries of the Pripyat, the boundary con- 
tinued beyond the USSR frontiers in the Brest region. 

The main advancement of the boundary to the south during the 
years 1947 to 1952 thus occurred in the region of the upper Dnieper. 
There the great northward curvature of the boundary line, so dis- 
tinct in 1928, totally disappeared. The boundary moved most exten- 
sively in the Volga region, between the Tsna and Volga, and in the 
Urals. Boundary advancement occurred in other regions also, but to 
a less extent. 

Available information about the northern boundary of the range 
in 1952 is sketchy. Evidently, however, the region of permanent 
habitation of the animal at that time reached the northern limits of 
the forests. 

In the 1940's and the commencement of the 1950's there were 
frequent visits by elk beyond the southern limit described above, 
often very far beyond. Along the Ural River visitations were recorded 
up to Ural'sk and Chapaev, in the steppes in the Dzhambeita region 
(east of Chapaev), in the steppes between Chapaev and Furmanov on 
the Great Uzen' River (50° N lat.), in the lower reaches of the Erus- 
lan, along the Khoper to Balashov, in the upper part of the Don 
basin in the Ertil' region (east of Voronezh), in the Lipets, Kulikov 
and Usmansk forests (left bank of Voronezh, north of the city of 
Voronezh), and even in the Boguchar region (slightly southeast of 
40° E long, and 50° N lat.). There were other distant visitations as 
well. In the Ukraine transgressions are known in the following 
districts: Poltav (almost up to Poltava), Sum, Chernigov, Kiev, 
Kamenets-Podol', Zhitomir, Volyn, and Roven.""^ 

The southern boundary of the region of fairly permanent habita- 
tion of elk in the Dnieper basin had already reached, or almost 
reached in 1952 the boundary of maximum range spread in 1880. 
Some individual visitations were recorded even beyond this line. 
The boundary lay even slightly more north (by a maximum of 100 
km) of the 1880 line at the source of the Oka and in the upper 
reaches of the Don and Voronezh. The two again coincided at the 
sources of the Tsna and Khoper. Later, along the Volga and between 
the Volga and the source of the Khoper (along the source of the 

^"Basedon data by D.N. Danilov, 1951; Stroganov, 1951; Martynov, 1951; Korneev, 
1952; Kirikov, 1952; Shvarts, Pavlinin and N.N. Danilov, 1951; Serzhanin, 1956; V.A. 
Popov and Lukin, 1949; Sludskii, 1950; Barabash-Nikiforovy 1957; and others. 



334 

Medveditsa), the 1952 boundary descended much farther south (by 
350 km) along the river than in 1880. The boundary lay that far 
south (Saratov region) only in the seventeenth and eighteenth cen- 
turies. Slightly west of the boundary line for the seventeenth and 
eighteenth centuries elk were only seen as transgressors (Balashov 
and Boguchar). 

The differences between the 1880 and 1952 boundaries are even 
more pronounced beyond the Volga: around 50° E long. (Sa'marsk 
Luka) the new boundary lay more to the south than before by 100 to 
150 km, and then around 55° E long, by 400 km or somewhat more. 
It should be borne in mind that the 1880 line described above refers 
mainly to permanent habitations, but partly also indicates visita- 
tions, while the new boundary line is confined only to fairly per- 
manent habitations. Transgressions at the end of the 1940's and in 
the first or second years of the 1950's, compared with the 1880 line, 
extended to distances (measured along meridians) of 150, 250, 350, 
400, or 500 km.''' 

From 1952 through 1958 the population and extent of coloniza- 
tion of elk continued to increase in most places in the European part 
of the USSR. Furthermore, in vast expanses along the southern part 
of the range conditions were created for supporting an extremely 
high density, bordering on overpopulation. As a result, not only 
were there deep and frequent transgressions beyond the range bound- 
ary but also some southward advancement of the region of perma- 
nent habitation. It was, however, not of such proportions as in the 
preceding decade since the entire region favorable for the habitation 
of elk to the south of the forest zone and in the forest-steppes was 
240 already occupied. The animal continued to occupy isolated forest 
masses along the southern edge of the forest-steppe region. In the 
Don basin such forests occurred along the Voronezh above the city of 
Voronezh (Lipets, Kulikov, and Usmansk— partly up to 1952), the 
Khoper (Borisogleb forest, Tellermanov grove; Khoper preserve in 
1952), and some other regions such as forests occurring southwest of 
Voronezh along the Sosna River, and Khrenov pine forests on the 
Bityuga River at Khrenovoe. Elk transgressions were noted in the 
Annen, Bobrov, Novo-Kalitven, Korotoyak, Ostrogozh, and Pavlov 
regions of Voronezh district. Here the animal was observed even in 
very small forest sections or even in open steppes far from forests, for 



^^*The 1947 boundary had already reached the 1880 position at some places. Along 
the Volga and in Trans-Volga it even crossed it (see map). 



335 

example in Dobrin and Bogucha regions (Barabash-Nikiforov, 
1957'"). 

Geographic range at commencement of latter half of 1950's and 
in 1958. In the north elk reached not only up to the northern 
boundary of the forest, but occupied the entire region of forest- 
tundra and led a normal and settled life up to the southern bound- 
ary of the tundra. This should thus be regarded as the northern 
boundary of the region of permanent habitation. Elk fully colonized 
forest-tundra after the 1950's, mainly from 1952 and 1953. Before this 
they were seen there only as transgressors or lived in small numbers 
and only at some places (V.Ya. Parovshchikov; Skrobov, 1958 and 
1958a). 

In summer large numbers of elk migrated deep into the open 
tundra and colonized it everywhere, reaching the coasts of the Bar- 
ents and Kara Seas. This occurred in all the tundras east of Kanin 
Peninsula. Arrivals in the tundra took place in spring (May-June) 
and at places resembled true migrations. In autumn (September-Oc- 
tober) a reverse movement led to wintering in the forest-tundra. '^^ 
While migrating, elk sometimes overcame barriers such as water 
bodies (rivers and gulfs), for example Korovinsk Bay on the right 
bank of the Pechora (Skrobov, 1958). The picture of seasonal range 
expansion on the Kola Peninsula is very clear: in the warm season 
animals occurred throughout the peninsula up to the sea coast, 
while in winter they remained only in the forest zone (R. Zakharov). 

In forest-tundra elk winter mainly along river valleys overgrown 
with osier, which serves as their food. Some animals winter even in 
the tundra zone, having been displaced from open tundra into the 
floodplains of much larger rivers (Pechora, Om', Pesha, and others), 
along which narrow strips of osier extend far into the north beyond 
forest-tundra. Thus elk winter even at the mouth of the Pechora. In 
winter they are sometimes found as transgressors even in open tun- 
dra (known up to 68° N lat.). 

On the Kanin Peninsula elk live year-round in forest-tundra in 



In the basin of the upper Don colonization was more successful than in other 
places. The natural conditions there are, of course, more favorable for such settlements 
(forest regions along the Don, Voronezh and Tsna). 

Even massive migrations have been reported (Skrobov, 1958). 
Regular seasonal migrations into the steppe, more so the tundra, are a new 
phenomenon in the biology of elk, at least over the last century. This ecological feature 
was acquired as a result of colonization. In the range a special region has been 
delineated, a region of summer habitation. This phenomenon is all the more 
significant since elk are essentially animals which lead a settled way of life. 



336 

small woods " between the Semzha (on the bank of Mezen' Bay) and 
the Nes'ya and winter even below Chizha where only willow thickets 
("ery") are available: in fact, they are so tall that they are never 
covered by snow. In summer elk are numerous in the open tundra 
north of the latitude of Chizha, wherever low osier beds exist. The 
animal reaches the northern limits of the peninsula (Cape Kanin) 
and the Kanin Kamen' mountain ridge in the north of the penin- 
sula. Along the northern slope of the Kamen' where tundra has a 
more severe Arctic climate and willows are absent, elk are not seen 
(E.P. Spangenberg; observations for 1956 and 1957). 
241 Since 1950, notably from 1955 to 1958, the number of elk asso- 
ciated with tundra, especially those remaining in it for wintering, 
has steadily increased. Evidently, a special population of elk has 
been formed right before our eyes in recent years, not only in the 
steppes (see below) but also in the tundra zone. 

Since 1951 the zone of distribution of elk to the south has also 
continued to enlarge. The southern boundary of the zone of perma- 
nent habitation has advanced and the region of visitation notably 
extended. This situation is explained by the fact that on leaving the 
zone of forest-steppes and steppes elk find only small sections conve- 
nient for permanent habitation. These are isolated, fairly large or 
sometimes even very small islands of forests. Visitations, however, 
extend even into the open steppes, sections of bottomland deciduous 
forests along rivers, small clearances, etc., i.e., into places where the 
animal cannot find satisfactory conditions for permanent habitation. 

It is also significant that, having colonized the steppes, elk have 
now penetrated into densely populated and wholly cultivated open 
expanses where they find no shelter whatsoever. This is due not only 
to the animal's relationship to man ("anthropophilous"), but also 
to wintering conditions. Thus in the Kur' district some elk spread 
far into the forestless eastern and southeastern parts in spring, 
summer, and autumn, raise their young there, and go back in winter 
into the northwestern part of the range where there are more forests 
(D. Starodubchenko). 

Transgressions are mainly irregular and aimless dispersals. 
Some of the animals die and some wander, probably for a long time, 
until they chance on some relatively favorable place in the south or 
return north. Thus two distinct modes of existence of the animal 
have recently been identified in the southern zone of the range as a 

Elk are often confined to such narrow and sparse forest and shrub "islets" that 
they cannot even hide in them on being frightened by man. Instead they run into the 
tundra and stay there until the source of disturbance has left the "islet". 



337 

result of movement farther soutfi: small individual islands of per- 
manent habitation along isolated forest masses or sections, and 
extensive areas of transgression with temporary, often very brief, 
residence. In these southern islands there are very few stray animals. 
Sometimes a few stray elk or a few dozen of them are sighted in a 
given region (Rovensk — about 15, Poltava — about 15, Kiev — about 
10, Chernikov — about 75, etc.; A.P. Korneev). In such regions all the 
intervening areas between individual islands are visited only occa- 
sionally. South of the peripheral points lies a zone in which elk are 
only visitors, but frequent ones; farther south lie some areas where 
elk penetrate very rarely. 

The southern limit and range boundary after 1955 and in 1958 
were as follows. In the far west the southern boundary covered 
sections of permanent habitation which have yet to be exactly 
defined. Evidently, commencing at the state boundary somewhere 
between Belovezh Forest and Brest it ran east, passing slightly north 
of Pinsk, or in the region of Pinsk, and left at the junction of the Styr 
with the Pripyat. 

From the mouth of the Styr the boundary runs upward along the 
river into the Roven district to the Stepan region (southwest of 
Sarna). From there it turns west through Klesov region of the 
242 same district (east of Sarna), runs in the same direction to the Olev 
region of Zhitomir district (Olev on Uborta), and later turns steeply 
southward into the Emil'chin region (Emil'chin at the source of the 
Uborta). From there the boundary line again runs west through the 
Lugin region to Korosten' (right bank of the upper reaches of the 
Uzh, tributary of the Pripyat^^'), and from there to the Razvazhev 
region of the Kiev district (lower reaches of the Teterev). Later the 
boundary line swings southwest toward Kiev. 

The western boundary of the region of transgressions in this part 

^'°The following outline of the southern boundary of elk distribution has been 
compiled on the basis of data (lists of regions of habitation, sometimes maps, etc.) 
kindly furnished by I.I. Barabash-Nikiforov (Voronezh), A.P. Korneev (Kiev — mainly 
data of the Central Board of Hunting, the Ukraine), Starodubchenko (Inspectorate of 
Hunting, Kur'), Mikhailov (Inspectorate of Hunting, Saratov), A.V. Fedosov and I.I. 
Ekimtsev (Inspectorate of Hunting, Bryansk), Sukhikh (Inspectorate of Hunting, 
Belgorod), Gusev (Inspectorate of Hunting, Rostov-on-Don), Vorshev (Inspectorate of 
Hunting, Stalingrad), and information from Severin, 1958; Alekseenko, 1958; 
Kompanets, 1958; Voloshko, 1958; Dolbanosov, 1957; and other data. 

^^'Only the administrative region as reported by the correspondents has been cited 
as habitation. As an explanation the geographic position in relation to the center of the 
region has been mentioned. 

^^^In the Zhitomir district there w^ere only about 20 settled elk in five regions (A.P. 
Korneev). 



338 

of the right bank of the Dnieper commences in the Kiverets region of 
the Volynsk district (Kiverets, northwest of Lutska), runs straight 
west through the Tsuman region of the same district, and later in 
the same direction into Gorodnits region of Zhitomir district 
(Gorodnits on Sluchi northwest of Novograd- Volynsk). From there 
it arced north-east through Barashev and Chernyakhov regions into 
Zhitomir. Later it ran west-northwest into the Makarov region of the 
Kiev district (east of Kiev) and southeast into the region of 
Rzhishchev. 

Two instances are known of extremely deep transgressions 
beyond this line — to the Gorodok region of the Khmel'ntsik district 
(about 50 km west-southwest of Proskurov) and into Solobkovets 
region of the same district (about 50 km southwest of Proskurov). 
These two interesting visitations in which elk almost reached the 
Dnestr show that the possibility is not excluded even now of the 
independent penetration of the animal into the Carpathian forest 
district where it lived at some time in the past. 

On the left bank of the Dnieper elk are dispersed extensively in 
the south in a large number of regions as a transgressor as well as a 
permanent inhabitant. This is mainly so in northern regions but 
also in some isolated regions far south. "^ 

The line limiting the southern territory in which there are sec- 
tions of elk inhabitation traverses the Dnieper at Kiev, later runs 
into the Borispol region (southeast of Kiev) and southeast to 
Pereyaslav-Khmel'nits. From there it turns east to the Chernukhin 
region of Poltava district (slightly east of Pripyat; Alekseenko, 1958), 
later southeast in the region of Dikanlka (north of Poltava), and 
straight north to the Gadyach region. From there, evidently bypass- 
ing Suma on the north, the boundary arced north and later east, 
passing into the Kur' district in the region south of Sudzha (Sudzhan 
region) or at the latitude of Suma. From there it runs east to the 
region of Staryi (possibly also Novyi) Oskol, then to Voronezh and 
southeast of Voronezh into Khrenov pine forest (KHrenovoe, slightly 
east of Bobrov). 

From Khrenovoe the boundary line runs east to the Khoper, 
descends along it to the south and, encompassing the Medveditsa 
basin, runs to Stalingrad. The southernmost point of elk habitation 
in the Stalingrad district is the Leshchev tree farm in the Lenin 



^^^The boundary given below is based on these southern points although the picture 
is not very clear north of it. Elk settled mainly in the south on the left bank region of the 
Kiev district and only transgressed in much of Chernigov (apart from the far north and 
northeastern parts) from 1955 to 1958 (data of A. P. Korneev). 



339 

region 40 to 50 km from Stalingrad. The largest number of elk 
inhabited the central part of the district, i.e., Frolov and Ilovlin 
regions (between the Volga, Don, and the lower reaches of the Med- 
veditsa). It is possible that the zone of permanent habitation of the 
animal also covered the far northern regions of Rostov district 
(Veshensk, Chertkov, and possibly others also). In any case this 
represents a place of "permanent encounters with elk" (A. Gusev). 

The region of visitations between the Dnieper and Volga to the 
south can be described as follows. From Rtishchev on the right bank 
of the Dnieper the boundary runs across the Dnieper in the Cherno- 
baev region of the Cherkassk district (slightly west of Zolotonosha), 
and from there following the general course of the Dnieper, to the 
Petrikov region of the Dnepropetrovsk district and toward Dne- 
propetrovsk. In this district there is on record a particularly deep 
243 transgression to the Novopokrov region on the right bank of the 
Dnieper northwest of Zaporozh'e (Voloshko, 1958). 

From Dnepropetrovsk the boundary runs east to Petropavlovka 
(at the mouth of the Byk River in Samara; Dolbonosov, 1957), then 
northeast to the Kremen region of Lugansk district (Kremenaya at 
the junction of the Krasnoi with the northern Donets), and from 
there southeast to the Staryi-Lugansk region (northeast of Lugansk 
beyond the northern Donets). Then it passes into the Rostov district 
at the Kundryuchya River (tributary of the northern Donets at its 
junction with the Don northeast of the city of Shakhta) and farther 
to the east into the Morozov region (south of the city of Morozov). 
From there, crossing the Tsimlyansk reservoir, the boundary passes 
into the Stalingrad district where instances of elk sightings in the 
steppes south of Stalingrad have been reported. Probably the bound- 
ary runs toward the Volga somewhere in the region of Cherni Yar. 
On the Volga itself transgressions of elk are known as far as Vladi- 
mirovka and Chernyi Yar. Possibly the animals descend from there 
along the floodplains of the river. 

In the region under consideration an extremely interesting 
encounter with elk occurred at the end of summer, and another in 
December, 1958, toward Sambek village located close to the coast of 
the Sea of Azov (Taganrog Gulf) between Taganrog and Chaltyr 
(eastern Taganrog, Neklinov region with its center at Pokrovskoe). 
Such instances provide convincing proof of the past habitation of 
elk through the steppes right up to the coast of the Sea of Azov and 
of contacts between southern Russian and Caucasian populations. 
Under conditions of light cultivation of the steppes, and in other 
natural situations (bottomland deciduous forests along the steppe 
rivers), it is natural that the animal had settled not only in that 



340 

region which is visited even at present, but also throughout the 
v\^hole area up to the southern seas. On the other hand current 
transgressions of elk tov^^ard Chernyi Yar serve as an additional 
proof in favor of the view that elk reached the Volga valley in the 
past and even the delta on the Caspian Sea. 

Thus the southernmost points of elk visitations lie roughly at 
49° N lat. (Solobkovets region), 48° N lat. (Novopokrovsk region 
and Chernyi Yar), and 47° 15' (Sambek; southernmost point in 
Europe and the Kazakhstan steppes). 

In the expanse between the Volga and Ural mountains and Ural 
River the boundary of the region of fairly permanent habitation of 
elk coincides with the Volga and runs north along the river to the 
mouth of the Great Irgiz. From there, along this river, it turns east 
and northeast and, in its upper reaches (already within the Kuiby- 
shev district), turned steeply southeast and, encompassing the 
Teplov region (north of Ural'sk), exited toward the Ural River. It is 
evident that along Ural valley elk are present quite regularly even 
below Ural'sk. Above it, they live permanently along the Ural valley 
throughout the Orenburg district. 

In the above region elk often emerge south into the steppe zones 
of Saratov district (recently, for example, Bezymensk, Krasnokut, 
Ershov, Ozinsk, and Klintsov regions) and also Stalingrad region. 
The southernmost points of visitations are the Urda region in the 
West Kazakhstan district (Urda, southeast of Lake El'ton), the steppe 
between Furmanov on the Great Uzen', and Chapaev (Lbishchensk) 
on the Ural at 50° N lat. and 50° E long., steppe slightly northeast of 
this place, Chapaev on Ural, and near the Ural in Taipak region at 
49°35' N lat. (Antonovo; Sludskii, 1953)."^' 

Thus only in the extreme northwest, north of 50° N lat., to the 
west of the Dnieper, does the boundary pass through the forest zone, 
where it was altered extensively. Throughout the rest of the range it 
lies in the zone of forest-steppe but mostly in the steppe zone; much 
of the region of permanent habitation lies in this zone. Finally, the 
zone of visitations not only adjoined the semidesert and desert zones 
but even covers a considerable part of them. 

Almost throughout its extension (apart from the far western sec- 
tion), the southern boundary of the range advanced far into the 
south compared with the 1880 boundary. On the right bank of the 
244 Dnieper by 1958 it had not yet reached the boundary of the seven- 
teenth and eighteenth centuries, but east of the Dnieper it penetrated 

^'""in some of the above places elk were reported somewhat before the year 1955 to 
1958. 



341 

deeper southward not only as a zone of transgressions, but partly 
(right bank of the Volga) also as a zone of permanent habitation. At 
some places the boundary of visitations has already approached, or 
even reached, the maximum possible reconstructed boundary. 

The recent (1930's to 1950's) map of elk colonization in the 
north, and especially in the south, represents a phenomenon of great 
importance, one influencing prevailing views about the ecological 
and geographic aspects of the species. This phenomenon is also of 
general biological interest (see below, colonization in Kazakhstan 
and western Siberia). 

Geographic Range in Western Siberia and Kazakhstan 

Reconstruction of the maximum range in this region over the his- 
toric period is possible only in a very general and partly tentative 
form because of the extremely scanty information available. In the 
north elk probably reached the forest boundary and the zone of its 
permanent habitation probably corresponded or was close to the 
present zone (Fig. 75). In the south it occupied the entire forest- 
steppe and the northern edge of the steppe zone. Between roughly 66 
to 67° and 71 to 72° E long., the range boundary formed a promi- 
nent projection southward, reaching the meridional course (upper) 
of the Ishim (up to 51°30' to 52° N lat.). In the east, roughly from 
Pavlodar to the latitude of Lake МагкакоГ, the boundary ran along 
the Irtysh, then turned east, and ran toward the state boundary (E.I. 
Strautman, 1953; map). The range covered all of the Altai up to the 
boundary with Mongolia and Pri-Altai steppes. Farther away the 
range crossed beyond the Yenisey. 

As in the case of the European part of the USSR, very significant 
positive and negative fluctuations of northern and southern bound- 
aries of the range also took place in the territory under description. 
Information about these fluctuations, except for the Recent period, 
is far less than that available for the European part of the USSR, and 
pertains almost exclusively to the nineteenth century. 

These movements, at least in the present century, have been 
occurring synchronously or fairly synchronously with westward 
movements. In any case, in the last decade the range of elk in western 
Siberia has enlarged notably both in the south and in the north. 
With respect to time and intensity, this enlargement has completely 
coincided with that occurring in the European part of the USSR. 
The northward movement of the animal in western Siberia and to 
the north and northeast in the European part of the USSR in partic- 
ular represents a single process. Evidently the range contraction in 



342 



45 50 55 60 65 70 75 80 85 90 



95 100 105 110 




,.4 -7 

2 ""I'niii 5 • s 

^^^^^ 3 wvwefTf» 6 '-"-<i—«-"9 

160 160 320 480k 

'■■■■' I I 



Siberia in the nineteenth century described below also coincided in 
time with the corresponding phenomenon in the west. 

Concomitant with enlargement of the region of permanent habi- 
tation, the number of visits beyond the limits of this zone has also 
increased greatly in the last decade. These transgressions have spread 
deeply in the north as well as in the south and elk have emerged not 
only into the forest-steppe, but also into the steppe, semidesert, and 
tundra zones. 

In the nineteenth century (in the first half and middle) the range 
of elk in western Siberia shrank strongly. Its area decreased to a 
minimum, especially in the north. Compared to the maximum pos- 
sible range (as at present) the northern boundary lay farther south- 
ward than in the west. Descending from the Urals, which it crossed 
around 60° N lat., the boundary crossed the Lozva roughly around 
61° N lat. From there it formed a northward promentory running 
initially northeast roughly up to 62°, later turned southeast, and 
emerged toward the source of the Konda River. Later the boundary 
turned toward the Ob', somewhat below the mouth of the" Irtysh, 
crossed the river, and ran for some time along its right bank not far 
from the valley. Slightly east of 75° it crossed to the left bank of the 
Ob', initially away from the river, later again coming close to it, and 
departed at the lower reaches of the Vasyugan. In the region of the 
mouth and lower reaches of the Ket' it again crossed to the right 
bank of the Ob'. From there, with a slightly southward arc,- the 
boundary ran along the right bank of the Ket' into the Yenisey 
roughly at 59°30' N lat. Beyond the Yenisey, though initially 



245 Fig. 75. Boundaries of distribution of elk (Alces alces L.) in Trans-Urals and western 

Siberia (scale in km). 

1 — northern boundary during maximum contraction of range in nineteenth century 
(from Nasimovich, 1955); 2— rough outlines of northern boundary in 1920's and ear- 
ly 1930's (from Formozov, 1946); 3 — boundaries of region of permanent habitation 
in 1920's and up to mid-1930's; between the two parts of the range and quite far in 
the north and south from the boundary line the animal appeared in those years as 
a visitor (from I.P. Laptev, 1958, with some modifications); 4 — northern boundary of 
region of fairly permanent habitation in 1940's and by 1951 to 1952 (V.G. Heptner); 
5 — southern boundary in these same years; 6 — northern boundary of region of per- 
manent habitation at commencement and middle of 1950's (1955; I.P. Laptev); 
7 — northern boundary of visitations to tundra (from LP. Laptev, 1958, with some ad- 
ditions); 8 — some points of deep transgressions in the north; 9 — boundaries in 
Kazakhstan (see Fig. 76 for southern boundary and transgressions southward). 

Note: Second boundary at the Ob', Ural, and Pechora appears drawn extremely deep 

into the north; in those years elk did not cross 65° N lat. and probably lived 

considerably more southward (62 to 63° N lat.). V.G. Heptner. 



344 

246 slightly ascending northward (but not reaching 60° N lat.), the 
boundary line inclined south slightly and passed eastward (Nasimo- 
vich, 1955; map). 

The southern boundary of the range at that tirne is not clear. 
However, judging from all the information available it lay quite far 
to the north and only very slightly more southward than the bound- 
ary shown below for the 1920's. 

In the latter half of the nineteenth century the range evidently 
enlarged slightly. It was obviously large at the commencement of the 
twentieth century. However, by the early 1920's it was in a state of 
extreme contraction. The reasons for this are not fully understood 
but there is no doubt that some of the important contributory factors 
were immoderate hunting and killing of elk. 

In its general outlines the northern boundary of the fairly per- 
manent habitation of elk in the 1920's and early 1930's represented a 
deep southward arc. It commenced at the Pechora somewhere 
around 65° N lat., intersected the Ob' around 63° N lat., emerged 
along its right bank toward the mouth or the lower course of the 
Vakh, and then passed eastward to the Yenisey, running slightly 
north of the uppermost course of Тут. At the Yenisey the boundary 
departed roughly around 61° N lat. and turned eastward, rising 
slightly north and emerging at the midreaches of the Podkamennaya 
Tunguska (Formozov, 1946). 

The above line reflects the northern boundary of only the zone of 
permanent habitation and that, too, schematically. The actual map 
of elk distribution in the 1920's and the first half of the 1930's is 
fairly complex. The range (zone of permanent habitation) is divided 
into two parts — western and eastern. The former was associated with 
the European part of the range and formed one single unit with it. 
Its northern boundary commenced at the Pechora somewhere in the 
region of the mouth of the Usa and proceeded toward the Ob', which 
was intersected at around 65° 30'. From there it turned southeast into 
the region of the source of the Kheta (left hand tributary of the 
Nadym) at roughly 64° N lat. and 71° E long, and later turned 
steeply west toward the Ob', encompassing the Kazym basin, and did 
not go within 80 km of the Ob'. Thus an eastward projection 
formed.'^' 



"^The range and boundaries described here and later are based on dataof I.P. Laptev 
(1958) for the period 1920 to 1935. There is, however, every reason to assume that in the 
1920's elk did not spread that far north along the Pechora and Ob'. At that time there 
was also no separately isolated section of permanent habitation of elk at the mouth of 
the Ob', as pointed out by the author (this section has not been marked). Along the Ob' 



345 

Farther away the boundary Hne turned straight south, leaving 
the Ob' roughly at 61 to 62° N lat., and descended along the Ob' up 
to the mouth of the Irtysh. Again intersecting the Ob' it ran along its 
left bank roughly up to the latitude of the mouth of the Тут and 
then grauually moved farther away from the Ob'. It intersected the 
lower reaches of the Vasyugan and ran southeast and south toward 
the source of the Om', initially in a slightly concave and later convex 
line. Having formed in this part of the Ob'-Irtysh interfluve region a 
large projection southeast, the range boundary became the southern 
boundary and turned west, initially through the upper reaches and 
later through the mouth of the Тага. Crossing the Irtysh there it 
later intersected the lowermost reaches of the Ishim and the lower 
reaches of the Tobol at the mouth of the Tavda, crossed the Tavda 
around 65° E long., and ran to the midreaches of the Tura. 

The boundaries of the eastern part of the range in the north and 
west are as follows. The northern boundary commenced beyond the 
Yenisey, slightly north of 60°, and running west intersected the 
Yenisey just above the mouth of the Podkamennaya Tunguska. On 
the left bank of the Yenisey it described a gentle arc and turned 
northwest, reaching its farthest point around 61°20' N lat. and 88° E 
long. The boundary turned south and ran sinuously in this general 
direction. It intersected 60° N lat. slightly east of the source of the 
247 Тут, intersected the Ket' at roughly the same longitude, crossed 
Chulym around Zyransk, and departed toward the upper reaches of 
the Tom' in the middle interfluve region of the Chulym and Tom'. 

In the expanse between these two regions of permanent habita- 
tion of elk lay a fairly broad region of 250 to 500 km where they were 
seen only as transgressors from one side or the other. Moreover, from 
both sides of the range elk moved considerable distances southward 
and very far northward (even up to tundra). Thus both zones of 
permanent habitation are linked by an extensive region in which elk 
transgressions are known. They were far from rare and a near-total 
exchange occurred between the two populations. 

From the 1930's (more from the middle or end of that decade) the 
elk population began to increase and its range also to enlarge. 



the boundary traversed evidently much more to the south, probably up to 63° N lat. It is 
evident that the line described (and the corresponding section in Laptev's map of 1958) 
pertained to theendof the period described by the author, i.e., 1935. In the 1930's theelk 
population had already begun to increase and the animal commenced to colonize, 
initially along the Urals and Ob'. The rest of the data of I. P. Laptev (1958) on range 
boundaries could be for the 1920's, i.e., theepochof maximum contraction of the range 
in the present century. 



346 

Initially poor, the population became extremely large in the 1940's, 
peaking in the 1950's. The two sections of the range, formerly sepa- 
rated, joined to form a continuous range and its boundaries began 
advancing south as well as north. At the very end of the 1940's and 
by 1950 to 1951 the northern boundary of the region of fairly per- 
manent habitation advanced north to a distance of 300 to 400 km 
and perhaps even more at some places. 

By 1950 to 1951 a definitive arcuate line had formed, which com- 
menced at the Ob' in the arctic circle at Salekhard (Obdorsk). From 
there it descended southeast quite steeply and intersected the upper 
reaches of the Nadym slightly north of 64° N lat. Running in the 
former direction, it proceeded farther toward the source of the Pur 
and, turning straight east, to the source of Taz. Turning northeast 
the boundary then ran toward the Yenisey evidently somewhere in 
the district of.Zyryanov (between 63 and 64° N lat.) and turned 
straight north along the Yenisey. It evidently proceeded along the 
right bank slightly away from the river. In the lower reaches of the 
Yenisey, at the latitude of Dudinka, it departed to the east (V.G. 
Heptner). 

Toward the mid-1950's (1955) the northern boundary shifted 
even farther. Intersecting the Ob' slightly below Salekhard, it ran 
southeast, intersecting 75° E long, not far from the coast of Tazov 
Bay and Nadym at 65° N lat. Then, taking an almost eastern direc- 
tion and only slightly inclining southward, the boundary inter- 
sected the Pur roughly at 64°40' N lat. and the Taz at 64°30'. It went 
off at 64° 15' N lat. toward Yenisey (LP. Laptev. 1958). 

On the whole, compared with the period of maximum range 
contraction in the nineteenth century, the northern boundary had 
advanced by a distance of 400 to 600 km or more by the mid-1950's. 
The northward advancement of the northern boundary continued 
during the latter half of the 1950's. 

In exactly the same manner as in the European part of the USSR 
elk penetrated far north beyond the above-described line into the 
forest- tundra and open tundra devoid of any forest vegetation. In this 
vast expanse (extending up to the sea coast) lying north of the bound- 
ary line visitations extended for a greater distance than in the west. 
They became more frequent and were on a larger scale in the 1940's 
and more so in the 1950's. At some places the summer movement of 
elk to tundra was a fairly regular phenomenon although it had not 
yet assumed the form seen in Bol'shezemel' and Malozemel' tundras 
in the west. 

On the whole the northern boundary of the zone of elk transgres- 



347 

sion lay in the Yamal and Gydan peninsulas, roughly at 71°00' to 
71°30' (LP. Laptev, 1958). 

The significant curvature of the northern boundary of elk, so 
sharply manifest in western Siberia and also in the Pechora basin, 
partly also in Trans-Yenisey Siberia, attracted attention in the 1920's 
and 1930's and even earlier (recent references include Yurgenson, 
1935; Podarevskii, 1936; Formozov, 1946; Nasimovich, 1955; LP. 
Laptev, 1958). At one time the view prevailed (Podarevskii, 1936; 
Formozov, 1946) that the absence of elk in the region of this curve 
was due to snow height there. In the Pechora basin, northwestern 
248 Siberia (Nadym, Pur, and Taz basins), and the basin of the lower 
Yenisey, the snow cover is maximum for the elk range as a whole 
(not less than 90 cm or so). 

The northern colonization of elk in the region of maximum 
snow described above proves that the snow cover should not be 
adduced as an explanation for the curve in the boundary. Hunting 
of the animal was of vital importance and deep snow only facilitated 
its extinction (Nasimovich, 1955). ^^^ That elk have not as yet per- 
manently colonized the forest zones in the basins of Nadym, Pur, and 
Taz, and along the left bank of the lower course of the Yenisey can 
only be a temporary phenomenon. If the dispersal process had not 
been artificially interrupted (by extinction), elk would still have 
continued to colonize these areas in the same manner as they did in 
the corresponding territories beyond the Yenisey (see below). At the 
same time it is possible, as sometimes assumed (LP. Laptev, 1958), 
that the penetration of the animal there was impeded or rendered 
difficult by the absence or inadequacy of some important food 
sources. 

The beginning of elk colonization in the south and the advance- 
ment of its southern boundary corresponded in time to the north- 
ward movement described earlier. It commenced evidently at the end 
of the 1920's and advanced perceptibly in the 1930's when the first of 
the visitations, sometimes over long distances, were noticed in the 
southern margin of the forest zone, the forest-steppe of western Sibe- 
ria, and some individual places even in northern Kazakhstan. This 
phenomenon assumed vast proportions in the 1940's. By the end of 
this decade in the south of western Siberia the elk had become a 
permanent inhabitant not only of the southern edges of the forest 
zone, but also of considerable parts of the forest-steppe. In the 1940's 



If not subjected to thoughtless persecution, and food sources are adequate, elk 
can survive in areas with a snow thickness of over one meter. 



348 

elk visited the steppes of western Siberia and the northern and east- 
ern periphery of Kazakhstan fairly extensively and became perma- 
nent inhabitants at some places. 

In the 1950's this process continued and elk colonized the steppe 
zone, although this period was evidently characterized not so much 
by an enlargement of the range, as by an increase in population and 
colonization of the animal at places where it was seen only as a 
transgressor in the previous decade. The natural limit of southward 
distribution attained its general features in the 1940's. 

The southern boundary of the region of permanent habitation of 
elk in western Siberia and Kazakhstan at the commencement of the 
1950's (by 1952) occupied the following position. Commencing in 
the west at Ural'sk, it proceeded along the Ural valley eastward to 
Orsk. From there, in a manner not precisely known, it rose north 
along the interfluve region of the upper Ural and the source of the 
Tobol roughly up to the latitude of Troitsk. Evidently, bypassing 
Troitsk to the north, or going through it, it turned steeply east 
toward the Tobol through the northern Kustanai district. Including 
the Vveden region (Vvedenka) within the range, the boundary 
turned sharply south toward the region south of Kustanai, and 
then encompassing islands of cut-over forests at Ara-Karagay to the 
east and southeast of Kustanai, turned east toward the Ishim 
through the region of Lake Ubogan."" There the boundary, run- 
ning in the same direction, traversed along the latitude 53°30' (at 
places probably even slightly more southward), including in the 
range all, or nearly all, of northern Kazakhstan in the east roughly 
up to 71° E long. From there the boundary inclined northward 
slightly and, passing north of Lake Seletytengiz, crossed the Irtysh 
around 52°30' to 53°00' N lat. From the Irtysh it proceeded in the 
form of a gende, slightly northward arc toward Lake Kulundinsk, 
passed it on the east, and running slightly east of 80° E long, turned 
to the Altai. 

It can be seen from the above brief description that by the early 
1950's (by 1952) elk had colonized not only the whole of the forest 
zone and the entire forest-steppe, but also a considerable part of the 
steppe zone. The region of transgressions by that time was much 
249 more extensive and the latter occurred so often at some places that 
a fairly permanent habitation or a tendency toward it may be 



Evidently, by that time elk already lived in Aman-Karagai pine forests roughly 
100 km south of Kustanai (north of Seminozernoe). There is no first-hand informa- 
tion on this, however. 



349 

assumed.^^^ In western Kazakhstan, along the Ural, elk intruded 
south up to 49°35' (Taipak region, West-Kazakhstan district) and, 
along the steppes east of the Ural, into the region of Dzhambeita and 
adjoining places (roughly 50° N lat. or slightly more). South of the 
Ural, between the river and parallel to Aktyubinsk and Orsk in the 
east, elk appeared in the steppe at several places. More southward, 
they reached the source of the Orsk (observed in the south up to 
49° 15' N lat.) even closer to the boundary of the semidesert. Farther 
to the east, after a fairly distinct break, frequent transgressions of elk 
have been recorded at several places to the north, west, and east of 
Kustanai. South of Kustanai elk intruded even as far as 52° N lat."'''' 

In the Kokchetav district, the one lying south of the North- 
Kazakhstan [district], numerous instances of transgression have been 
recorded throughout almost all of the district to the east of 71° E 
long. It is possible that in the northwestern part of the district elk by 
then had begun to colonize or tended to do so. South of this part of 
the Kokchetav district instances of transgression were known in the 
northern part of Akmolinsk [district] to 52° N lat. and even slightly 
more to the south, to Atbasar, i.e., to the Ishim between 68 and 70° E 
long. No transgressions were reported in this period between 71 and 
73° E long. Elk occurred between 74° E long, and the Irtysh only in 
the far north of Kazakhstan; farther south a transgression was 
reported at 74° E long., roughly as far as 52°30' N lat. 

Along the right bank of the Irtysh elk normally lived in the 
region of Lake Chany and south of it, and along the Pri-Altai 
steppes with their ribbons of pine forests. Transgressions from there 
in the direction of the Irtysh and to the Irtysh itself were known at 
several places throughout the stretch of the river to the south almost 
up to Usk-Kamenogorsk.^'*'' In the very south they probably came 
from the Altai. 



Evidently the situation along the Ural valley below Ural'sk (Chapaev and 
Taipak regions). 

From the region of Kustanai and Ubogan and from Ara-Karagai forest at 
Kustanai, which elk had already colonized, to this place for not less than 150 km, 
about 100 km is an open steppe with only tiny lakes up to Aman-Karagai pine forest 
(at Semiozernoe) and the rest again steppe. Two transgressions have been recorded 
there, one in 1939 and another in 1940. 

The dispersal and dynamics of the elk range in western Siberia and Kazakhstan, 
in spite of innumerable articles and minor notes referenced in the text, are based 
on data compiled by Middendorff, 1851, 1867, 1869, and 1874; Yurgenson, 1935; 
Kulagin, 1932; Formozov, 1946; Yanushevich and Blagoveshchenskii, 1952; and others, 
but mainly based on the data and compilations of Sludskii, 1953 (Kazakhstan); 
Nasimovich, 1955; and LP. Laptev, 1958. 



350 

Thus in the early 1950's elk were present in all the places in 
which they were known in the eighteenth century. 

In some of these regions, reckoned above as places of transgres- 
sion, the animal during this period was already resident and 
multiplying. 

Unlike the European part of the USSR, no significant changes 
occurred after the 1950's in the disposition of the southern boundary 
of the general zone of elk distribution in the southern part of western 
Siberia and Kazakhstan. The animal evidently reached the natural 
limit even before this period. In any case, in 1958, apart from some 
particularly deep transgressions, elk were generally encountered 
within the boundaries of 1952. Possibly some expansion of the 
region of transgressions took place west of Tobol (western Kusta- 
nai), but on the other hand there was some retreat to the west of the 
district boundary, north of the upper reaches of the Ishim (A.A. 
Sludskii). 

It is evident, however, that the population of the animal and its 
density increased in the southern zone of the range. As a result, there 
were some changes in the nature of residence of the animal in some 
individual places in the range under discussion: in places where elk 
were formerly transgressors they began living permanently or 
almost so, and in places where transgressions were rare, the same 
251 became more regular, the boundary of permanent habitation had 
slightly advanced southward by 1958 but positive data about this 
phenomenon are not available. 

The increase in elk population along the southern margin of the 
range is also borne out by deeper, farther southward transgressions 
in the 1950's. In recent years, before 1958 and in that year, elk 
intruded into the region southwest of the source of the Ilek (in 
Tamir region), the region between Chelkar settlement on the rail- 
way line and Mt. Irgiz, the Dzhetygarin region of Kustanai district 
(north of the source of the Tobol), the Ulutau mountains, the region 
between Lake Tengiz and the Ishim, the mountains in the region of 
Bayansk-Aul, and the mountains in the region of Karkaralinsk (A.A. 
Sludskii). These points lie in the zone of steppes, at Temir in the 
semidesert, and at Chelkar even in the desert zone. 

In all the maps on elk colonization in the steppes (in Kazakh- 
stan) certain areas stand out prominently. In the west movement 
proceeded from one side along the Ural valley, and from another 
side, also in relation to the Ural valley, from the Orsk region in 

^"'This is probably associated with the natural changes occurring in the northern 
parts of Kazakhstan in recent years and represents a special local phenomenon. 



351 

Aktyubinsk. In the central part of Kazakhstan the broad front of 
colonizing animals moved into the area from the zone adjoining the 
Tobol in the west up to roughly 71° in the east. In almost all cases 
migration was associated with river valleys, trees and shrubs in bot- 
tomland deciduous forests, clearances in steppes and forest sections, 
and pine groves in steppes and in small elevations, and rarely along 
rushes by lakes. This part of Kazakhstan is particularly favorable in 
this purely biotopic context. Sometimes the animals moved south 
and into open steppe and, more interesting, even penetrated the 
semidesert and true deserts. 

It should be emphasized that this front, especially between 67° 
and 71° E lat., is moving in the direction of those places where elk in 
Kazakhstan existed in the eighteenth century, and thus the restora- 
tion of the former range proceeds very intensely. Furthermore 
although there are no direct references to it, elk formerly lived also 
in the forests north, west, and east of Kustanai and also in the pine 
forests of Aman-Karagai 100 km south of the town. 

In western and central Kazakhstan advancement was from the 
north. In a third of the zone, along the Irtysh, colonization pro- 
ceeded probably mainly from the east from the Pri-Altai steppes 
with their ribbons of pine forests and the Kulundin steppe and, to a 
lesser extent, from the north from the region of Lake Chany and 
Omsk. 

Very little information is available about elk distribution in the 
Altai. In the period of peak range expansion in the historic period 
elk occupied, as shown before, the whole of this montane area up to 
the state boundary, apart from places like the Chuisk steppe which 
are unsuitable for survival. In the south the range evidently extended 
into the Narym range and probably also to the Kurchum; the bound- 
ary traversed Lake Магка-КоГ or slightly north of it. This was 
possibly the situation by the early nineteenth century. Subsequently 
elk range in the Altai shrank drastically. It continued to connact at 
the end of the last and at the commencement of this century when, 
generally speaking, this process ceased in western Siberia. By the 
1930's elk had practically disappeared in much of the Altai and were 
preserved only in the easternmost and northeastern parts by the 
middle of that decade. There elk occupied the Chulyshman basin 
and part of the basin of the Bashkaus (a tributary of the Chulysh- 
man) and the region of the upper courses of Bol'shoi and Malyi 
Abakan and the source of the Lebed'. Slightly to the west they 
occurred in the zone adjoining Lake Telets from the west and in the 
source of the right-hand tributary of the Katun, i.e., Kodrin (Dmi- 
triev, 1938; V.G. Heptner). 



353 

The boundary outlined above may be regarded as corresponding 
to the state of utmost reduction of the range. Later, but before the 
recent period, ahhough it enlarged there is almost no positive sup- 
porting information. Range expansion and population increase evi- 
dently occurred not only in the eastern section described above, but 
also with the arrival of animals from the north and their coloniza- 
tion of the foothills, histances are known, for example, of the 
appearance of elk around Biisk (Kuchin, 1958). It is possible that elk 
transgressions in the upper Irtysh (toward Ust-Kamenogorsk — see 
above) were associated with their increasing population and range 
expansion in the Altai. 

252 Geographic Range in Trans-Yenisey, Siberia, aiid the 
Far East 

Reconstruction of the elk range east of Yenisey in the period of its 
maximum development in the historic period is extremely difficult 
because of scanty information. Data are available to show that in 
the Taimyr, elk reached 71 to 72° N lat. (Middendorff, 1869), 
approached very close to the Kolyma estuary (Filippovka River; 
Wrangel, 1841), and were widely dispersed in extreme northeastern 
Siberia. In general, however, with the exception of some places, the 
boimdaries of the maximum range in the past could be regarded as 
closely corresponding to those that existed in the middle and latter 
half of the 1950's. 

As in western Siberia during the same period (commencement 
and middle of the last century), a contraction of the range took place 
in central Siberia also. This process evidently did not cover the 
eastern part, i.e., the Lena basin and farther east. Boundary fluctua- 
tions did take place there, for example in the Kolyma, but they were 



250 Fig. 76. Boundaries of distribution of elk {A Ices alces L.) in the Trans-Volga, south of 
western Siberia and in Kazakhstan (scale in km). 

1 — boundary of Kazakhstan; 2 — southern boundary of forest zone (taiga); 3 — northern 
boundary of steppe zone (between lines 2 and 3 — forest steppe); 4 — northern boundary 
of semi-desert and desert zone (between lines 3 and 4 — steppe zone); 5 — southern 
boundary of zone of permanent habitation of elk in the early 1950's (by 1950; based on 
data of Siudskii, 1953 and I. P. Laptev, 1958, with some modifications and additions); 
6 — southern boundary of region of elk transgressions in the 1930'sand 1940's and early 
1950's (by 1952), and some points of transgressions within it; 7 — some especially distant 
transgressions of elk before 1952; 8 — boundary of zone of elk distribution in 1958; 9 — 
especially deep transgressions of elk at the end of 1958 (nos. 8 and 9 from A. A. Siudskii); 
10 — zone of elk habitation in Kazakhstan in the past. Boundaries of natural zones 
schematic. V.G. Heptner. 



354 

evidently exceptions and their magnitude small. 

During the period of extreme recession the northern boundary of 
the range commenced at the Yenisey roughly at 59° 30' N lat. and 
turning east ran to the uppermost course of the Podkamennaya 
Tunguska. In this expanse the boundary formed a gentle arc slightly 
curved northward. However, it did not run along the above river and 
did not reach 60° N lat. From that place on the Podkamennaya 
Tunguska the boundary line made a steep loop and encompassed 
from the south the interfluve region of the sources of the Podkamen- 
naya and Nizhnyaya Tunguska, turned north along the upper 
reaches of the Nizhnyaya Tunguska (slightly east of it), and crossed 
the river in the region of 63° N lat. From there, slightly inclining 
west, it ran to the upper reaches of the Kotuya (Nasimovich, 1955). 
Farther away in the east the northern boundary even at that time lay 
far in the north, probably along the forest boundary. 

In the present period (middle and latter half of the 1950's) the 
northern boundary in this territory commenced at Lake Pyasino 
(east of Dudinka on Yenisey; V.N. Skalon). In a manner not clearly 
understood, but probably along the northern boundary of forest- 
tundra, i.e., along 71 to 72° N lat., it proceeded evidently to the 
mouth of the Khatanga or the lowermost reaches of this river and 
later into the lower reaches of the Anabar. It intersected the Anabar 
at 72° N lat. (slightly south of Syskylakh) and ran in the form of a 
slightly irregular line to Olenek around 71°30' N lat. From there it 
proceeded to the Lena, running toward it at Kyusyur or slightly 
north of it (about 71° N lat.).^^^ 

East of the Lena, corresponding to the shrinkage of the range in 
the tundia, the boundary ran southeast, proceeding on 70° N lat., 
and later again rose northeast to the Omoloi slightly south of 71° N 
lat. From there it ran to the Yana, intersecting it at Kazachego or 
slightly north of it (71° N lat.). On the Khorma the boundary went 
only slightly southward, but formed a slightly southward bend 
between 138 and 144° E long., not reaching 70° N lat. On the Indi- 
girka the range boundary went slightly south of the Chokurdakh, 
intersecting the river roughly at 70°30' N lat. From there the bound- 
ary ran southward. It intersected 150° E long, at 70° N lat., Alazeya 
at 69°30',Chukoch'ya around 69° N lat. and the Kolyma at Nizhni 
Kolymsk or slightly farther north (around 68°30'). At this latitude, 



^^^It is possible that along the Lena, as also along some other rivers in Europe and 
Asia, elk extended far into the north. Instances of transgressions are known even in 1928 
to Cape Bekov in the Lena delta at 72°30' N lat. (Kolyushev, 1936). 



355 

descending slightly farther, it ran into the basin of Bol'shoi and 
Malyi Anyuya beyond Yakutia. 

This boundary in Yakutia (i.e., between the Anabar and Kolyma) 
traversed along the southern boundary of the tundra, partly through 
forest-tundra. As a result of the general increase in numbers, com- 
mencing from 1954, fairly regular seasonal (summer) dispersal of elk 
into tundra was observed, especially east of the Yana, the animals 
reaching several places along the sea coast. The dispersal evidently 
occurred mainly along rivers but also in open tundra. Thus, 
between 140 and 142° E long, elk emerged up to 72°. In the region of 
the lower reaches of the Khorma up to its mouth (72°), along the 
Indigirka up to its delta (almost 72°), along the Alazeya, and in 
253 tundras adjoining it from the east, the animals reached the sea 
around 71° N lat.^'^ 

The range is known poorly to the east of the Kolyma in far 
northeastern Siberia. In this region, essentially devoid, or almost 
devoid, of forests, conditions for survival of elk are hardly favorable. 
The animal is forced to confine itself to river valleys, mainly in 
willow thickets, and at places poplar groves. As a result, not only is 
its population very low but also the boundary itself bears a complex 
outline and is extremely unstable. It is very difficult to demarcate the 
region of fairly permanent habitation from that of transgressions of 
individual animals. Moreover, the available information pertains to 
very early periods, i.e., the 1920's and 1930's. 

From the above place in Kolyma the boundary ran east, encom- 
passing Malyi and Bol'shoi Anyuya basins, and later turned south, 
covering the upper reaches of the Anadyr or at least the region of the 
left hand tributaries of the upper Anadyr — the Yablon and Yeropol. 
It is possible that elk dispersed (transgressions?) downward along 
the river roughly to Markovo. From there the boundary ran south 
into the source of Penzhina and, leaving the Taiganos Peninsula 
outside the range, ran toward the Okhotsk coast in the northern part 
of Gizhigin Bay. Formerly elk normally reached Markovo along the 
bottomland deciduous forests of the Anadyr and, in the remote 
period, evidently even up to the lower reaches of the Anadyr or even 
the sea coast (Kanchalan River, flowing into the Anadyr estuary, 
Amerikanskaya Koshka).^"*"* 

In the distant past (seventeenth and eighteenth centuries) elk 



Range boundary in Yakutia is based on unpublished data of D. Ivanov (Yakutia). 

^'''' According to data published by Portenko, 1941. According to Abramov, 1954 

(map), the boundary ran along the midportion of the Bol'shoi Anyuya, upper course of 

the Malyi Anyuya, and through the sources of the Yablon, Yeropol, and Penzhina. 



356 

were distributed more widely and probably were encountered along 
the bottomland deciduous forests and other rivers of the northeast- 
ern extremity of Siberia. Instances are known of elk remains from 
the ocean shore at Cape Schmidt (Nordenskiold, 1870; cited by Nasi- 
movich, 1955). Deep transgressions are known even more recently, 
for example in 1936, on the Bol'shoi Baranikh River around 69° 30' 
N lat. and 166°20' E long, at a distance of 120 to 150 km north of the 
forest region (Portenko, 1941). It is possible that in the past and 
possibly in the recent period, from the upper reaches of the Penzhina 
the boundary ran along the river to the coast of Penzhina Bay, or 
Penzhina was frequented by wanderers. 

From Gizhigin Bay the boundary ran along the sea coast south- 
ward to the Ussuri region; elk were absent in Kamchatka, the 
Shantar Islands, and Sakhalin. Along the coast of the Ussuri 
region the boundary ran south roughly up to the mouth of the 
Amgu River (around 46° N lat.). From there it withdrew from the 
coast into the mountains and extended in the form of a narrow 
projection along the more elevated parts of Sikhote-Alin up to the 
source of the Iman (around 44°45' N lat.; Kaplanov, 1948). In the 
middle of the last century, and also much later, it extended along 
Sikhote-Alin up to the upper courses of Ussuri and Suifun, i.e., up 
to 43° 30' N lat. (Przewalski, 1870). In the region of the mouth of the 
Imansk the range boundary intersected the Ussuri, exiting in the 
west into China. 

In the expanse from the Ussuri to the Altai the elk range cur- 
rently crosses through the southern boundary of the state or runs 
along it, as for example in the Tannu-Ola range. The range boundary 
deviates from the state boundary somewhat significantly only in the 
eastern and western Trans-Baikal steppes, which it encircles from 
the north. 

This is evidently the situation also in some other places, but the 
curvature of the boundary line to the north is less significant and 
hardly associated with topographic features (extremely high moun- 
tains for example). On the whole the southern boundary of elk is 
known only very generally. 

Geographic Range outside the Soviet Lhiion 

At present""' the American range of elk is considerably reduced 

^"'According to Abramov (1954) the boundary ran toward the Okhotsk coast only in 
the region of Tauisk Bay; it extended northward away from the sea along the divide of 
the Kolyma range. 

^''^See the section on western Siberia for elk distribution in the Altai. 

^"'866 the description of the genus for the reconstructed range. 



357 

although it preserves in general the outlines of a zonal range, 
254 extending from Alaska to Nova Scotia. Unlike the range of Ameri- 
can wapiti, of which only three prominent and some tiny sections 
have remained, it is not only still very extensive but also forms a 
single block (Burt, 1952; Seton, 1909-1910). As in the USSR, in 
America also elk became better adapted than deer to living in the 
proximity of man. Some periods are known in recent times during 
which elk advanced slightly into the south and at places into the north. 
However, there has been no significant increase in range, at least not 
to the extent seen in the Soviet Union. 

In Europe, in much of its natural range, i.e., in western and 
central Europe, elk became extinct even around the tenth century. 
Up to the eighteenth and nineteenth centuries elk survived only in 
the far eastern parts (not including the USSR) — in Saxony until 
1746, Galicia until 1760, Silesia until 1776, and Polish Pri-Baltics 
until 1830. In Rumania (Carpathians) elk evidently died out in the 
eighteenth century. In the twentieth century and before the Second 
World War some (at places considerable) elk'were present in eastern 
Poland. At present in the German Democratic Republic and Federal 
Republic of Germany elk are altogether absent and present in 
Poland only in the northern and northeastern parts. They are com- 
mon in Finland, Sweden, and Norway. On the Scandinavian Penin- 
sula the elk population has increased significantly and the animals 
have advanced extensively into the south in recent years and wholly 
colonized the peninsula up to its southern extremity. ^"^^ 

In Asia, outside the USSR, elk live in Mongolia and northeast 
China. In Mongolia they occur in the Hobsogol region in the east 
and in the west up to the State boundary of the USSR, along the left 
bank of the Egin-Gol and Selenga in the east up to 105° and in the 
north up to the frontiers and Hentei. Here elk live in the south to 
Bogdo-Ula (slightly southeast of Ulan-Bator) and in the west to the 
Tola, almost up to its confluence with the Orhon. The Khentei 
section of the range joins with the Soviet section in the expanse 
between 108 and 1 1 1° E long, and forms a projection of the Siberian 
range southwest (Bannikov, 1954). 

In northeast China elk live along the northern part of the Great 
Khingan, slightly penetrating even into the easternmost corner of 
Mongolia at Il'Khuri-Alin and the Little Khingan. Evidently elk 
also inhabit the northern parts of the eastern Manchurian montane 
region, but information regarding this is inadequate. (V.H.) 

In recent years instances are known of the appearance of elk on Zealand Island in 
Denmark. The animals swam through Sund Strait, which is about 10 km wide at its 
narrowest point. 



358 

Geographic Variation 

Geographic variation in elk has been well studied and the litera- 
ture is quite substantial. Yet views on the number of species and 
geographic races vary considerably. ¥oy the entire range of the spe- 
cies, including America, two species with seven subspecies have been 
proposed (Flerov, 1931 and 1934). Of these, both species and three of 
the seven subspecies occur in the Soviet Union. According to 
another view (Buturlin, 1934) a single species of elk is acknowledged 
for both the Old and New World and four subspecies for the Old 
World and the USSR. Finally, it was recently suggested (Flerov, 
1952) that there is only one species of elk with a single form for 
Europe and western Siberia, one for the Caucasus, and one for all of 
the Trans-Yenisey part of the range and the whole of the American 
range. 

Elk are fairly mobile and a considerable population mixing, 
especially in certain periods, has occurred, hindering the emergence 
of intraspecific geographic differentiation. Nevertheless, it does exist 
and there is no doubt that the recent view about the existence of only 
two forms of elk (not counting the Caucasian) is erroneous. In the 
Old World and the USSR there are four fairly well defined forms, 
possibly even six (see below). In the New World there are evidently 
only two subspecies. The subject calls for further study with ade- 
quate material. The following forms are present in the USSR. 
255 1. European elk, A. a. alces Linnaeus, 1758 (syn. coronatus, euro- 
paeus, machlis, antiquorum, palmatus, jubata, typicus, uralensis, 
meridionalis, and tymensis). Medium to fairly large animals but not 
the largest. Body length of bulls up to 270 cm, height at withers up 
to 216 cm, and weight up to 500 kg or more (in exceptional cases up 
to 620 kg). Total length of skull of bulls 550 to 580 mm and length of 
rostral part (from anterior edge of orbit to tip of premaxillae) 329 to 
347 mm. Antlers with fairly well developed, sometimes well- 
developed palmate portions with a span of 135 cm; weight (without 
skull) up to 20 kg. Palatine processes of premaxillae in front of 
vomer narrow and form a groovelike depression; total width less 
than, or equal to, vertical diameter of infraorbital foramen. Nasal 
processes of premaxillae not constricted upward but enlarged like a 
spatula with a blunt terminus (Flerov, 1952). Color of animal de- 
scribed earlier (legs light-colored). 

Found in the European part of the USSR, Urals, in the east 
roughly to the Yenisey, western Siberia, and Altai. Outside the 
USSR occurs in Europe. 

This form is not completely homogeneous throughout its exten- 




359 



Fig. 77. Rostral portion of skull of elk (from Flerov, 1952). 

Top — European and western Siberian {Alces alces L.); Bottom — Yakutian {Alces alces 

pfizenmayeri Zuk.). 

sive range. Animals inhabiting western Siberia are significantly 
larger, heavier, and have more powerful antlers with a distinctly 
developed palmate portion. Those in the European part of the 
USSR, especially from the more western parts, have a body length of 
250 to 270 cm, height at withers of 175 to 185 cm, and weigh up to 
480 kg (usually 320 to 400 kg). Western Siberian animals attain the 
maximum of the above limits in size, weight, and antlers. 

Before the Revolution differences were largely associated with 
the fact that in the European part, in view of excessive hunting, 
killing of the best males, and some other reasons, the range of these 
animals narrowed down and at places even disappeared totally (in 
the west). In western Siberia the population was preserved under 
normal conditions and the animals could attain full development. 
There is, however, no doubt that differences were also due to natural 
factors, the above conditions only accentuating them and rendering 
them sharper in the last decade before the Revolution. Under present 
optimal conditions for animals in the European part of the USSR 
elk are completely protected. 

The taxonomic status of western Siberian forms requires a study 
based on new material from Siberia and Ural. The name uralensis 
Matschie or tymensis Zukowski may prove suitable for western Sib- 
erian elk. 

2. Caucasian elk, A. a. caucasicus Verstschagin, 1955*. Small size. 

*Spelling of author's name in taxonomic divisions is sometimes at variance with 
spelling in text and bibliography because the Israeli orthography has been used in this 
translation — General Editor. 



360 

Skull length 538 mm (one animal) and length of upper tooth row 
136 to 138 mm. Skull similar to European type. Nasal processes of 
premaxillae enlarged at ends, sometimes reaching nasals. Facial part 
in rostral region more intensely contracted than in northern elk. 
Palmate portion of antlers very poorly developed. Coloration and 
body dimensions not known. 

Found in forests of premontane plains in northern Caucasus, 
bottomland deciduous forests, and reed thickets on the Kuban and 
Terek. Evidently also occurred in the lower reaches of the Don and 
probably the Kolkhid lowlands. Extinct by early nineteenth century. 
Outside the USSR not known, but probably penetrated adjoining 
parts of Turkey by way of Batumi. 

3. East Siberian or Yakutian elk, A. a. pfizenmayen Zukowski, 
1910 (syn. yakutskensis, angusticephalus, 7 bedfordiae; the names 
americanus and gigas are used in Soviet literature for this form). 
256 Larger than western elk. Body length up to 300 cm, even more in 
exceptional cases; height at shoulders 225 to 240 cm; weight 570 kg 
or even 620 to 655 kg. Skull length in males up to 585 to 620 mm and 
width of zygoma up to 240 mm. 

Antlers very large and massive, usually with large and broad 
palmate section and large number of short tines, including large, 
stunted, and bifurcate anterior tines separated from palmate section. 
Antler span up to 159 cm, width of palmate section up to 60 cm, and 
circumference of rosette up to 42 cm; antler weight may considerably 
exceed 20 kg. Palatine processes of premaxillae in skull in front of 
vomer broad and flat and do not form a groovelike depression; their 
total width exceeds vertical diameter of infraorbital foramen. Nasal 
processes of premaxillae constricted and ends honed, wedgelike, and 
not enlarged like a spatula. 

Coloration same as in European elk but with slightly greater 
admixture of gray tones in winter and slightly darker, sometimes 
altogether black limbs, almost indistinguishable in color from trunk 
(Flerov, 1952). Dewlap usually larger than in western forms. 

Found in Siberia east of the Yenisey and in the Far East except 
the Amur and Ussuri region (i.e., in the south evidently to the Stano- 
voi range). Outside the USSR occurs in the northern periphery of 
Mongolia (Pri-Kosogol, Khangai, and Hentei). 

Elk in this territory represent a special form, well distinguished 
not only in body size and some color characteristics (less specific) but 
also craniologically. These features (especially the craniological) 
draw the eastern Siberia elk rather close to the American moose 
which, from the viewpoint of zoogeography, is quite understandable 
and natural. In fact an attempt has recently been made (Flerov, 1952) 



361 

to place all elk from the entire Trans-Yenisey Asiatic part of the 
USSR (including the Ussuri region) under the form americanus 
described from the northeast USA, grouping them with moose of 
North America, including Alaska. This viewpoint, insofar as the 
taxonomy of eastern Siberia elk is concerned, cannot be accepted. ^''^ 

Throughout the extensive range of its distribution this form is 
not really identical. Northeast Siberia i.e., the KolymaTndigirka 
basin and forests of the Anadyr basin are colonized by particularly 
large-sized animals, the largest of all elk seen in the USSR (maxi- 
mum among Russian elk), with extraordinarily large antlers. It has 
been suggested (Buturlin, 1934) that this form constitutes a distinct 
race proximate to, or identical with, the large elk of Alaska {gigas 
Miller). An independent subspecific status of elk in this region 
appears justifiable but the subject requires further study. 

4. Ussuri elk, A. a. cameloides Miline-Edwards, 1867 (syn, ? bed- 
fordiae and bedfordi; the name americanus is also applied to this 
form). Smallest form of elk, much smaller than the European A. a. 
alces from the European part of the USSR. Compared with western 
Siberia elk, it is not only smaller but stands lower on its legs, has a 
relatively larger head, and a lighter, more slender body. Body 
volume less. Body length of males (five animals) 250 to 276 cm (M 
256), height at withers 172 to 195 cm (M 179), height at croup 170 to 
180 cm (M 174), chest 173 to 180 cm (M 176), tail length 10 to 13 cm, 
and length of ears 22 to 24 cm. Maximum recorded weight of bull 
400 kg (Kaplanov, 1948) but usually does not exceed 320 kg 
(Abramov, 1954); average weight probably only slightly more than 
200 kg. 

Skull measurements are generally smaller than those of Euro- 
pean elk and the rostral region in the Ussuri form is relatively short- 
ened. In structural features skull corresponds to that oi A. a. 
pfizenmayeri (eastern Siberia type). Color uniformly dark while legs 
usually light -colored. 
257 Antlers usually small and slender, without palmate portion, 
branched, dichotomously divided, circular in section, with large 
tines, usually symmetrical; in general antlers resemble those of deer. 
Number of tines in fully grown animals varies from two to four on 
each antler, reaching five in rare cases. Antler span 80 to 100 cm and 
weight 5 to б kg. Exceptionally (less than 2% cases), formation of 
rudimentary palmate section observed but area does not exceed 100 
cm' (in western Siberia elk, 1,000 to even 2,500 cm^; Kaplanov, 
1948). 

Neither is an interrelation with American forms, at least the peripheral 
americanus and gigas acceptable (see below). 



362 

Found in the Ussuri and Amur regions in the north up to the 
Stanovoi range and the Lena watershed. In the west found roughly 
up to 123° E long. In the north along the Stanovoi range and near 
the sea much larger individuals occur and animals with palmate 
antlers, either intermediate to Yakutian form or representatives of 
latter forms have been reported. Elk of Sikhote-Alin are particularly 
typical. 

Ouside the USSR occurs in northeastern China in the east, 
including the Great Khingan. References to a find in Khentei are 
erroneous as the animals living there are A. a. pfizenmayeri. 

The form under description has been extremely well character- 
ized and is sharply distinguished from all remaining forms of the 
species, especially the eastern Siberia A. a. pfizenmayeri, which has 
large antlers with palmate section. Combining the two into a single 
form (Flerov, 1952) cannot be justified. 

The antlers of this form are of particular interest from the view- 
point of evolution of deer antlers. In other populations, especially in 
the western European part of the USSR, antlers without palmate 
sections are also seen, but in older animals. This feature in usually 
characteristic of underdevelopment and often associated with deteri- 
oration of a population or old age. The foregoing notwithstanding, 
such antlers are less massive and differ in structure from those of 
Ussuri elk. 

The features of Ussuri elk are associated with their past distribu- 
tion in the remote Manchurian-Chinese region. Evidently many 
primitive features are preserved in this form. 




Fig. 78. Head of Ussuri elk (Alces alces cameloides M.-E.) (from Flerov, 1952). 



363 

In some parts of the range falling outside the USSR, i.e., in 
Eurasia, the animals resemble those in the Soviet Union. Several 
subspecies have been described from America but actually there are 
only two: A. a. gigas Miller, 1899 in Alaska, Yukon, and parts of 
British Columbia (largest of all the forms of the species with antler 
span running up to 200 cm) and A. a. americanus Clint., 1822 in the 
remaining parts of the American range. (V.H.) 

Biology 

Population. The following numbers of elk were recently counted in 
the territory of RSFSR (N.V. Eliseev): 

1950 — 266,100 1956 — 353,000 
1954 — 310,000 1960 — 480,000. 

Supplementing these figures with data for Belorussia, Lithuanian 
Soviet Socialist Republic, Kazakhstan, and other republics of the 
Soviet Union, the elk population in USSR territory could number 
500,000 animals. Data for the RSFSR are based on reports received 
from the Directorates of Hunting of various districts, territories, and 
258 republics, which often determine elk population by counts (aerial 
and ground surveys), replies to questionnaires, or, rarely, general 
impressions. In some regions these figures are obviously exagger- 
ated; it would therefore be more prudent to put the total strength of 
elk in the USSR at 400,000."° 

There is no doubt that in the USSR the elk population at present 
is larger than at any time in the last 150 to 200 years. This is the 
result of a rise in numbers almost everywhere and tremendous 
expansion of the range, or more precisely, restoration of the former 
range over the past 20 to 30 years (see above). The elk population 
from 1928 to 1947 alone in some regions of the European part of the 
USSR rose on the average 10 to 15 times (Danilov, 1951; Dolgushin, 
1953; Sludskii, 1950 and 1953; Nasimovich, 1955). 

Data on elk populations in different regions have been given 
selectively because of the difficulty in obtaining complete informa- 
tion and in several cases because of its absence. In Belorussia in 1927 
about 150 elk were counted (Fedyushin, 1929), in 1936—380 
(Kulagin, 1940), and in 1952—600 to 800 (Serzhanin, 1955). Most elk 

^^"Between 200,000 to 300,000 elk have been counted in Canada (early 1950's), 
11,600 to 17,900 in the USA (1937 to 1948), 30,000 in Alaska (1946) (Peterson, 1955), 
20,000 in Finland (1957), and 120,000 in Sweden. An average of 21,800 elk were caught 
annually in Sweden in the early 1950's (Sainio, 1956; and others). 



364 




Fig. 79. Elk around Moscow, Zaveta Ilich station, Northern Railway. August, 1959. 
Photograph by T.V. Koshkina. 



259 



were in the Poles'e, Pinsk, and Viteb districts. In the northernmost 
Ukraine only a small number live a settled life; most are transgres- 
sors from Belorussia or RSFSR. The population on the whole is 
around 100. In the Baltic republics the maximum number of elk 
occurs in the Lithuanian Soviet Socialist Republic where there were 
300 in 1939, 90 in 1948, and 400 in 1955 (Ivanauskas, 1957). 

In the European part of RSFSR elk density per unit area is 
maximum in the central part of the range where mixed forests, and 
coniferous forests slightly more north, predominate. In the latter half 
of the 1940's the mean reserves of elk per 1 ,000 hectares of [mixed] forest 
area were 0.8 to 1 .0, in the subzone of northern coniferous forests 0.6 to 
0.7, and in the southern parts of the range where deciduous trees 
predominate and pine groves are scattered only 0.3 to 0.4 (Danilov, 
1949). 

The following examples provide some idea about the distribu- 
tion of elk in the territory of Russian Soviet Federal Socialist Repub- 
lic and partly also their population dynamics. In the Smolensk 
district in the winter of 1936 to 1937 there were 500 elk, which rose to 
around 1,500 by 1950 (Grave et al., 1951) and to 3,600 by 1954. In the 
Moscow district 200 animals were counted in 1929 (Yurgenson, 
1935), 1,550 Ш 1944, 2,030 in 1946 (Danilov, 1947), and 9,000 in 1956 
(Aleksandrova, 1957). In the Leningrad district in 1957 there were 
about 10,000 elk and in Pskov over 20,000 (N.V. Eliseev). In the 



365 

Vladimir district in 1954 there were 2,260 elk (Aleksandrova, 1957); 
in the Gorkii 1,720 in 1946 (Danilov, 1950); 4,250 in the Perm in 1941 
(Danilov, 1949) later reaching 10,000 (Kuklin, 1946); in the Sverdlov 
in 1957 there were over 20,000 elk (N.V. Eliseev); in the Tatar 
Autonomous Soviet Socialist Republic in 1929 a total of 83 (Apisov, 
1930), which by 1950 had risen to 1,318 and by 1954 to 2,400 (Aspisov, 
1954). Some 25,000 elk were counted in the Arkhangel'sk district in 
1951, which rose to 30,000 to 33,000 in 1954 (Semenov, 1957) but fell 
to about 25,000 in 1957 (NV. Eliseev). The elk population in the Komi 
Autonomous Soviet Socialist Republic is quite considerable and 
slightly less in the Karelian Autonomous Soviet Socialist Republic. 

In the latter half of the 1940's the average number of elk per 1,000 
hectares of forest area in Siberia was placed at 0.4 to 0.5 (Danilov, 
1949), i.e., only slightly more per unit area than in the southern parts 
of the range in the European part of the RSFSR. In 1957 in the Amur 
territory 12,000 elk were counted, 4,000 in Primor'e, 10,000 in 
Khabarovsk, over 8,000 in the Tuva Autonomous District,^^^ and over 
25,000 in the Yakutsk Autonomous Soviet Socialist Republic (N.V. 
Eliseev). In 1941 there were 7,500 in Tyumen and Kemerov districts 
together (Danilov, 1949). 

The elk population continues to rise in the territory of Russian 
Soviet Federal Socialist Republic. By the early 1950's elk had become 
quite common in northern Kazakhstan; their occurrence was record- 
ed in nine districts (Sludskii, 1953). Recently, a slight reduction in 
elk population has been noted in Kazakhstan. 

Habitat. Within the Soviet Union elk inhabit almost every zone 
of the forest and are even found in forest-tundra, being confined 
there in winter to small spruce-larch forest islands in protected mon- 
tane valleys (Polar Urals), willow thickets in swales and along the banks 
of rivers, and also birch and aspen groves (Dolgushin, 1953). At places 
in summer elk emerge into the open tundra, sometimes several hun- 
dred kilometers away from closed forest boundaries; some penetrate 
right up to the shores of northern seas. In winter the animals migrate 
southward into forest-tundra (Leble, 1953; Makridin, 1955; Gromov 
and Rapoport, 1938; Skrobov, 1958; and others). 

Elk are fairly common in forest-steppes but confined mainly to 
islands of forests (deciduous^^^ and pine) or marshy forests. Along 

"' About 4,000 elk were counted at the end of the 1940's in the Tuva Autonomous 
District (Skalon and Shikin, 1950). 

" In forest-steppes of the European part of the USSR elk are often encountered 
in oak forests with good reforestation, in young aspen groves (in winter), and also 
in plantations containing elm, maple, etc. 



366 

river valleys small numbers of elk penetrate the steppe zone where 
they live along the banks of rivers and lakes overgrown with willow 
and appear in summer in sedge marshes, among reeds, and even in 
the open steppe 100 to 150 km from the southern boundary of per- 
manent habitation (Sludskii, 1950; Danilov, 1951; and others). In 
Stalingrad region elk have been seen at the end of summer in the 
semidesert zone along banks of canals overgrown with willow, 
poplar, and maple (Stroganov, 1951). 

Thus the prevailing view that elk are purely taiga animals is 
incorrect. This view arose in the period when elk became nearly 
extinct in the central regions of the European part of the USSR. The 
fact that this animal reached the banks of Azov and Black Seas a few 
centuries earlier through river valleys was subsequently overlooked. 

In addition to lowland forests, elk were earlier widely distributed 
even in montane taiga. Right up to the end of the eighteenth or early 
nineteenth century they also inhabited hornbeam and beech forests 
of the northern Caucasus and, in the even more distant past, of the 
western Trans-Caucasus (Vereshchagin, 1949; and others). In mon- 
260 tane taiga elk prefer a relatively less rugged, gentle relief, and avoid 
steep slopes; broad valleys with abundant water sources or swamps 
represent particularly favorable spots. The vertical range of distribu- 
tion is broadest in the eastern Altai and the Sayan ranges where the 
animal is found in summer right from the foothills to the upper 
forest boundaries (1,800 to 2,000 m). In the Altai they are also found 
on balds, especially at places where there are small lakes with 
swampy banks overgrown with willow and dwarf birch (Dmitriev, 
1938). Thus in the Altai elk are found in summer roughly 2,200 to 
2,400 m above sea level. They also occur above timberline in summer 
in the Sikhote-Alin mountains in tall grass meadows (up to 1,600 m; 
Kaplanov, 1948). In Lapland preserve about 5% of all elk counted 
occurred in montane tundras where the animal is confined to willow 
thickets (Semenov-Tyan-Shanskii, 1948). The forest boundary there 
runs at a height of about 350 m and elk ascend mountain slopes 
roughly to a height of 500 m. 

Over a significant stretch of their range in the forest zone 
summer and winter habitats of elk differ sharply, but the animals 
visit during the greater part of the year willows along rivers, some 
types of deciduous forests with a tall herbaceous cover and fairly 
well-developed undergrowth, and also marshes overgrown with wil- 
low, dwarf birch, and pine. 

In the warm period of the year, especially in summer, preferred 
habitats are those which have rich aquatic vegetation along water 
юurces, whose banks are overgrown with willow, shrubs, or even a 



367 

real forest. Among the habitats used regularly in summer are 
swampy alder groves and marshes (often overgrown with willow, 
birch shrubs or pine), especially if they are interspersed with small 
dry ridges overgrown with forests or adjoin large forests. In addition 
to water sources where elk feed on aquatic vegetation, other impor- 
tant summer habitats are mixed and deciduous forests with well- 
developed and fairly tall herbaceous vegetation. Other important 
habitats are young, burned-over and felled sections with luxuriant 
grass cover, areas with shoots of deciduous trees, and areas with 
abundant willow herbs [fireweed, Epilobium] (an excellent food for 
elk). The greater the variety of food sources, the more favorable the 
area for elk. In most dense forest masses with a uniform composition 
of trees elk are never numerous either in summer or winter. 

Good shelter, offering protection from blood-sucking flies, is 
very important in the life of elk (see "Daily Activity"). Where elk are 
threatened by man in summer, they like to hide during the day in 
dense thickets of small forests, swampy alder thickets, or in dense 
coniferous saplings where their detection is difficult. 

The winter distribution of elk is determined mainly by the avail- 
ability of tree and shrub foods and also the thickness and density of 
snow cover in different biotopes in the latter half of winter. As far as 
possible, elk avoid sections with a snow cover of over 70 to 80 cm; in 
mountains and rocky sections shaded slopes with very loose snow 
are preferred. At places where elk are hunted the availability of 
shelters for protection from enemies are also of considerable 
importance. 

Elk often use the banks of water sources as wintering sites, espe- 
cially of rivers overgrown with willow, birch (in the north), and 
other deciduous species, growing burned-over and felled clearings, 
deciduous scrub woods (aspen and birch), and mixed and coniferous 
forests with dense undergrowths or numerous young trees. Elk often 
winter in pine forests rich in young trees (wintering of elk in such 
forests in the European part of the USSR has been observed from the 
farthest southern part of the range up to the Kola Peninsula inclu- 
sive) and in boreal forests (fir and spruce) with an abundant under- 
growth of rowan [mountain ash], birch, goat willow, and others 
(Baltic, Leningrad, and Arkhangel'sk districts, Komi Autonomous 
Soviet Socialist Republic, the Sayans, Bureinskiy mountain range, 
Sikhote-Alin). Nut pine forests are of least importance; elk are found 
in them (eastern Siberia and Sikhote-Alin) only if other coniferous 
261 and deciduous species are also present in large numbers. Larch 
(Larix) forests quite often serve as a habitat for elk in summer as well 
as in winter. Mixed forests are usually preferred to purely coniferous 



368 



^ 







Fig. 80. Group of elk in Senchinsk marshes in upper course of Om' River, western 
Siberia. January, 1957. Photograph by K.T. Yurlov. 

or deciduous. The only exception in this respect is young pine 
groves where large numbers of elk generally winter. 

Elk are found in winter even in marshes, but only when these are 
extensively overgrown with willow and birch or, in the north, dwarf 
birch. Elk avoid marshy sections with ice crusts under the snow. The 
environs of marshes represent favorite haunts of elk in early spring 
in western Siberia (Dem'yanka River) since snow clears earliest in 
such areas (Kaplanov, 1935). 

Depending on natural and other conditions, a particular habitat 
acquires special importance for elk. For example, in several regions 
of northeastern Siberia elk gather in winter in willow thickets along 
river banks (Zonov, 1931; Mikhel', 1937; Portenko, 1941; and others) 
or even congregate in overgrown burned clearings along the slopes 
of ridges where willow is abundant (Verkhoyansk badlands*; S.P. 

*Literally, folded lands— Sci. Ed. 



369 

Naumov). The winter habitat of elk in the basin of the Chul'chi 
River (eastern Ahai) at a height of 1,400 to 1,700 m is highly typical; 
there, the animals are mainly confined to broad mountain valleys 
overgrown with round-leaved birch (Betula rotundifolia), in the 
proximity of the edge of deciduous or mixed forests with an abund- 
ant undergrowth of shrub birch {Betula humilis). Aspen, mountain 
ash, and birch, common in much of the winter habitat of elk, are 
almost absent in the composition of these local forests (Dmitriev, 
1938). 

No less characteristic is the habitat-related distribution of elk in 
Sikhote-Alin. In summer elk are confined mainly to coniferous 
forests (nut pine-spruce-deciduous and larch) but also move into the 
floodplains of rivers, marshes, the upper portions of mountains in 
tall grass meadows among plantations of rock birch [Betula 
ermani), and also in grassy-spruce forests. In winter the maximum 
number of elk occurs in open larch and spruce-larch forests and in 
boreal forests with deciduous underbrush (Kaplanov, 1948). 

Often males are distributed in very different habitats from those 
of females with calves. For example, males locate in summer mainly 
in the upper courses of Altai rivers near timberline and on balds, 
while females with calves are common along lower slopes (river 
valleys and forest lakes) (Dmitriev, 1938). Only some adult males 
and females without calves remain in the watershed regions of 
Pechora-Ilych preserve, which is covered by pine groves, in the latter 
half of winter. The majority of elk and all females with young are 
confined to river valleys with willow thickets, since on the ice of 
262 rivers there is less snow than in the watershed regions (Teplov and 
Teplov, 1947). In Lapland preserve females with calves winter in the 
least snowy sections; only males and females without calves locate in 
areas where the snow is greater (Semenov-Tyan-Shanskii, 1948). The 
wintering sites of males in Sikhote-Alin are usually located at higher 
elevations than those of females (Kaplanov, 1948). 

Food. Within the Soviet Union elk, according to Danilov (1958), 
feed on 149 genera of angiosperms (1 10 genera of herbaceous plants 
and 39 genera of Qees and shrubs). They also eat 5 or 6 genera of 
gymnosperms (pine, juniper, yew, and others), no less than 5 genera 
of pteridophytes (except for horsetail, others are seldom eaten), 3 to 4 
genera of lichens (mostly epiphytic and rarely the terrestrial Clado- 
nia), 11 genera of fungi, several genera of algae (laminaria, Con- 
jerva, and others), and sometimes even mosses. In all elk utilize as 
food up to 175 genera of plants, representing no less than 250 spe- 
cies. This subject has been variously studied in different regions but 
by and large data are scanty. 



370 




Fig. 81. Winter habitat of elk in a felled section of Pri-Ob' pine forests. Novosibirsk 
region. January, 1957. Photograph by K.T. Yurlov. 



263 



In Mordov preserve observations on two pasturing domesticated 
elk established that in the summer-antumn period they feed on 133 
species of plants, of which 61 constituted principal food items 
(Shaposhnikov, 1951). In Priok-Terras preserve (Serpukhov region 
of Moscow district) elk in summer eat no less than 62 plant species 
(Aleksandrova and Krasovskii, 1957). In Lapland preserve elk use no 
less than 47 plant species, of which 14 constitute primary food items 
(Semenov-Tyan-Shanskii, 1948). In Oka preserve (Ryazan' distiict) 
in the summer-autumn period alone elk eat 46 species (Borodin, 
1940), and in Sikhote-Alin 43 species^^^ (in summer — 32, of which 6 
are primary; in winter — 28, of which 11 are primary; Kaplanov, 
1948). For Belorussia, on the whole, 38 species have been identified 
(Sablina, 1955; Serzhanin, 1955), and for Zhiguli in summer 33 spe- 
cies (Zharkov, 1954). Information is much more scanty for other 
regions of the country. 

Seasonal variations in food are distinct among elk. In summer 
the main food is represented by herbaceous plants, among which 
many are aquatic, grow around water bodies, and in general flourish 



"'According to Abramov (1954) Far East elk consume over 90 plant species but the 
author has actually identified less than 30, all of which were previously identified by 
Kaplanov (1948). 



371 

in moist habitats. Among other herbaceous plants elk avidly con- 
sume relatively tall plants (willow herbs [fireweed] are a particular 
favorite), sticky-leaved spirea, buckbean, buttercup, sweetflag, horse- 
tail, Nuphar, water lities (not only the green portions but also the 
rootstock), and many others (calla, cat-tail, Sagittaria, Alisma, 
Potentilla, loosestrife, dandelion, sorrel, some Umbelliferae, etc.). 
Information on these plants comes from a relatively small number 
of places in the range. 

Over much of the range (Kola Peninsula, Arkhangel'sk region, 
Komi Autonomous Soviet Socialist Republic, Rybinsk reservoir 
region, Altai, and Yakutia) sedge is utilized quite well but mainly in 
spring and early summer. The role of grasses, especially low ones, 
not only in the Soviet Union but also in America (Peterson, 1955), is 
not generally important in the diet of elk. Taller grasses at times are 
consumed in large quantities in the southern parts of the range 
(Tula abatis [stockade], Mordov preserve, and Zhiguli) and at places 
in the north (Rybinsk reservoir). For example, in the stomach of an 
elk caught in Tula abatis in mid- August, grasses predominated: reed 
grass, bent grass, meadow grass, hair grass, etc. (Likhachev, 1957). 
Elk in Zhiguli grazing sites consume in August about 90% giant 
fescue plants (Zharkov, 1954). In Mordov preserve five types of 
grasses serve as primary food items (Shaposhnikov, 1951). In Priok- 
Terras preserve sedges, leguminous plants, and grasses are almost 
untouched by elk (Aleksandrova and Krasovskii, 1957). 

On the whole elk are quite adaptable and probably less fastidious 
in their choice of food^^"* than some other ungulates. Elk can live and 
reproduce in habitats even in the absence of such important summer 
foods as aquatic plants (Zhiguli). In this case the absence of some 
food plants is evidently compensated by the presence of others. For 
example, in Zhiguli those consumed avidly from May through 
August are Aegopodium, Crepis, and Bupleurum (Zharkov, 1954). 
These plants are usually not known as food of elk in other parts of 
the range.^^^ In the diet of Ussuri elk 25 to 30% of the plants utilized 
as food belong to the Manchurian floral complex (Kaplanov, 1948; 
Abramov, 1954); in other parts of the range these varieties of plants 
are absent. With migration from one area to another, the food com- 



Basedon observations of grazing domesticated elk, Shaposhnikov (1951) came to 
the conclusion that the food composition largely depended on the degree of plant 
abundance. This conclusion is extremely debatable and is not in conformity with the 
winter observations of the same author (see below). 

^^'in Mordov preserxe Aegopodium is considered a secondary food (Shaposhnikov, 
1951). 



372 

position of elk also changes notably (Shaposhnikov, 1951^ Likha- 
chev, 1957). 

In summer trees and shrubs, mainly leaves and green shoots, are 
commonly used as food to a much lesser extent than in winter. Quite 
often elk do not eat the branches completely, tearing off only the 
leaves by running the branch through their mouth. Among conifers 
elk eat only young pine shoots in summer (Dem'yanka River in 
western Siberia; Rybinsk reservoir). Elk eat leaves of aspen,^^^ moun- 
tain ash, some species of willow, and buckthorn best (mainly in 
autumn, together with fruits). They also consume well bird cherry, 
oak, and some species of birch, at least in some periods of the vegeta- 
tive season. Moreover, they consume well in some regions for at least 
part of the summer period, leaves or shoots of maple (Zhiguli and 
Sikhote-Alin), raspberry (Belorussia, Mordov preserve), linden 
264 (Mordov preserve), ash (Belorussia), Euonymus (Zhiguli), elm, dog 
rose, blackthorn, spirea (Buzuluk pine forest), etc. It is interesting 
that in Priok-Terras preserve elk are not at all attracted to oak, 
Euonymus, and linden in summer. There are references for other 
regions where Euonymus, linden, and other species are poorly or 
not at all consumed by elk. 

Herbaceous plants and trees are generally consumed throughout 
summer to various degrees, but aquatic plants are consumed well 
throughout the warm period from the time they begin to grow well 
(end of May to early June) until the end of September or even 
October inclusive. Terrestrial herbaceous plants are consumed in a 
definite sequence, since they grow at different times, and the animals 
prefer tender green portions and parts which at a given time are 
more nutritious: the first shoots, tender leaves, and stalks in spring 
and early summer, and flowering shoots in the latter half of 
summer, i.e., the upper portions of stalks with bud, ovaries, and 
flowers (Shaposhnikov, 1951; Zharkov, 1954). 

In Zhiguli in May herbaceous plants account for about 79% of the 
entire intake and shoots with tender leaves, sharp-leaved maple, and 
bird cherry the remaining 21%. Leaves oi Aegopodium, Bupleurum, 
grasses, and chervils [cow parsley] constitute the basic food. In 
June-July the intake ratio of herbaceous food to tree food remains 
the same (80 to 20%), but along with Aegopodium and Bupleurum, 
leaves of Crepis and spirea are also well consumed. In August the 
proportion of tree foods increases to 40% (up to 90% terminal shoots 
of oak and 81% willow). Flowering shoots of over 10 species of plants 

The stomach of an elk killed in August in the basin of the Northern Dvina was 
filled with aspen leaves (Yurgenson, 1935). 



373 

are eaten well — up to 88% Bupleurum, 83% fireweed, 74% Crepis, and 
50% broad-leaved bluebells, etc. (Zharkov, 1954). 

Similar shifts in the composition of the food intake of elk have 
also been traced in Priok-Terras and Mordov preserves and in many 
other regions. In Mordov preserve over most of the summer a 
marked diversity in the food consumed is evident and no perceptible 
preference for particular species discernible. In spring and autumn 
the food composition is less diverse (in August about 50 plant spe- 
cies, in September-October about 20, and in November, 6) and a few 
plant species predominate during these periods of the year (Sha- 
poshnikov, 1951). 

In Lapland preserve the main food in spring (end of May to first 
half of June) is represented by fresh leaves of willow, birch, and 
mountain ash. From July considerable quantities of fireweed, spi- 
rea, sedge, marsh horsetail, buckbean, Nuphar, etc. are consumed. 
In September herbaceous vegetation predominates and in October 
mainly tree food. The summer food intake is quite extensive, the 
items widely dispersed, and only some plant parts, mainly the upper 
portions of herbaceous plants, nibbled (Semenov-Tyan-Shanskii, 
1948). 

Secondary foods of elk in the warm period, not consumed univer- 
sally but fairly rarely, include algae, ferns (in Mordov preserve 
bracken and ostrich feather ferns are well consumed by tame elk), 
tops of reeds, and most berries. On the other hand stalks and leaves 
of berry shrubs (bilberry, mountain cranberry, and blueberry) are 
quite well consumed. In September and October some acquire 
nearly as much importance as primary foods (Belorussia, southern 
Karelia, Arkhangel'sk region, Komi Autonomous Soviet Socialist 
Republic, and Sikhote-Alin). In Mordov preserve in the last ten days 
of October leaves and stalks of mountain cranberry have comprised 
42.5% of the food intake and bilberry 15.2%. The proportion of 
mountain cranberry in November was 42.3% (Shaposhnikov, 1951). 
265 In Mordov preserve tame young elk at the end of summer and in 
autumn consume nearly 15 species of mushrooms (including fly 
agaric). In the first ten days of October about 20% of the food intake 
is accounted for by mushrooms (Shaposhnikov, 1951). Mushrooms 
are avidly consumed by elk in the Pechora basin (Turov, 1953) and 
in the southern part of the Far East (Abramov, 1954). Similar obser- 
vations for other regions are absent; some researchers (Cherkasov, 
1884) contend that elk do not eat mushrooms. Only a few have 
recorded the consumption of mushrooms in Canada (Peterson, 
1955). The intake of mushrooms by elk is quite well known in 
Sweden (Skuncke, 1949). Evidently elk respond 4ifferently to mush- 



374 

rooms in different parts of the range. Herbaceous plants consumed 
by elk comprise some poisonous ones and some of medicinal value 
(in Priok-Terras preserve lily-of-the-valley is eaten well). Buckbean 
and calla possess anthelminthic properties (Rykovskii, 1957). 

In winter tree and shrub vegetation, mainly branches, and most 
needles of coniferous plants represent the main food of elk. In the 
early, less snowy period of winter, and sometimes throughout winter 
in the southern region of the range, deciduous varieties play the 
main role, while the importance of conifers perceptibly increases in 
the latter half of winter.^" Contrary to the highly nutritious food to 
which elk turn in summer, tree food is essentially supportive in 
winter. Elk survive on tree food for six to eight months, for a shorter 
period in the southern part of the range, and a longer period in the 
north where winter is more prolonged. Elk are better adapted to 
such foods than the overwhelming majority of ungulates. The Even- 
kian allegorical names for elk are quite apt: "moota" (tree eater) and 
"shektata" (willow eater). For superstitious reasons Evenkians 
while hunting avoid using the common word for elk, "toka" 
(Podkamennaya Tunguska River; Suslov, 1927). 

The range of trees and shrubs consumed by elk in winter is 
extremely broad — no less than 50 genera'^^ — of which roughly one- 
fourth form the main diet in one region or another. In some parts of 
the range relatively few play a major role: in the northern part and 
in typical taiga regions not more than 4 or 5 genera, in the central 
regions of the European part of the USSR (Moscow and others) 6 to 
8, and in the same area but in the zone of broad-leaved forests and 
forest-steppes 10 to 12. In Sikhote-Alin elk readily consume 8 to 10 
species of tree species. 

In a given region not more than half of the total number of trees 
available constitute primary food at a given time. This is especially 
true of regions in which the latter half of winter is extremely snowy 
and access to low shrubs is greatly hindered. For example, in 
Lapland preserve in the latter half of winter elk rarely consume 
266 small willows and junipers which are normally their favorites 
(Fig. 82), but feed mainly on branches of birch and pine needles 
(Semenov-Tyan-Shanskii, 1948). 

Among the tree and shrub species of importance to elk in winter 
as primary food in much of the range in the USSR are willow, 

^"At places where the population density of elk is high and deciduous forests few, 
pine is quite often intensely attacked even from early winter. 

It is not precisely known which species (highly numerous) of willow, birch, and 
pine are consumed by elk. Still, the intake of different species of a given genus varies 
notably. 



375 



XI-XI 



XII- 



IV-V 



1UU 
80 


; 




= 




















60 

40 

20 




- 


ЬИ. 


^ 


^ 


,-■■ 


^ 




П 




\ 






П 


\ 






Pine 




Birch 








111! 


Junipe 


r 








Willow 







265 Fig. 82. Ratio between intake by elk of preferred twigs (willow and juniper) and 
principal (birch and pine) food in relation to height of snow cover. Lapland 
preserve (from Semenov-Tyan-Shanskii, 1948). 

Note: Roman numerals refer to calendar months. 

pine'^^ (especially prominent is the role of saplings and young trees 
up to the age of 15 to 25 years), aspen, and mountain ash. Birch is 
eaten equally extensively but less avidly; however, at places where 
aspen and mountain ash are few (northern parts of the range) birch 
is considerably more important than the other two species. In a very 
limited part of the range species important as primary food are 
juniper (some taiga regions of the European part of the USSR), fir 
(Pechora basin, Urals, Irkutsk region, and Amur zone), bird cherry 
(many regions of the European part of the USSR and Sikhote-Alin), 
maple, oak, and willow (southern parts of the range within the 
European part of the USSR; in Sikhote-Alin maple also). Buckthorn 
and warty-bark spindle tree are well eaten (central and southern 
parts of elk range within the European part of the USSR) but not 
abundant everywhere; hence they do not constitute primary food 
items though highly preferred by elk. In this same category belong 
currants (a few varieties) and raspberry, which are eaten well but 
usually limited and, moreover, quickly covered with snow. In a few 
regions alder, linden, and hazel nuts are consumed satisfactorily, but 
rarely. Species such as poplar, honeysuckle, acacia, dwarf birch, rho- 
dodendron, and yew (rare everywhere) are usually not significant for 
elk. Almost everywhere spruce is eschewed; its consumption by elk 
usually serves as a reliable index of an unfavorable food situation. 



^^'in America pine is hardly touched (Peterson,1955). 

^*°Fir Abies balsamea (absent in the USSR) is an important winter food of 
American moose in eastern Canada (Peterson, 1955). 



376 

The uneven intake (sometimes from excellent to total rejection) 
of a given plant species in different parts of tfie range may be due to 
the differences in their chemical nature and morphology, as well as 
the degree of abundance and accessibility of other, more preferred 
foods. In forests adjoining Oka preserve, at places where there are 
many elk, pine is usually the main food (100%). Young trees are 
often completely destroyed, 76% have broken tops, and up to 20% 
wither away. In forests where elk are few and food more readily 
accessible pine is eaten much more rarely as elk prefer aspen, moun- 
tain ash, gray willow, and buckthorn over pine (L.P. Borodin). Pine 
and birch represent main and bulk food; although compulsory they 
are nowhere preferred as food by elk in most regions. 

A good idea of the food selectivity of elk is provided by compara- 
tive data on tree species consumed by them and the availability of 
those species in forests (Table 1). 

In winter elk damage shoots of deciduous trees and shrubs (up to 
20 to 30 cm long and 5 to 12 mm thick at the bite point); in the case 
of coniferous plants, needles and rarely shoots are consumed. Dur- 
ing below-freezing weather, bark is poorly eaten and, for example, 
in the north at temperatures under -2° С bark is almost untouched 
(Fig. 83). In the southern parts of the range where winter is very mild 
and thaws frequent, bark on the southern side of trunks is warmed 
by insolation even on frosty days and thus consumed throughout 
winter in large quantities. For example, in the Tula abatis, accord- 
ing to Likhachev (1939), bark is the most important constituent of 
the winter diet of elk. The importance of bark is particularly dis- 
cernible in transitional seasons of the year (spring and especially 
autumn); its consumption is far higher in those months than at any 
other time of the year. In summer elk rarely touch bark. Sometimes 
they eat only shoots (of birch, hazel nut, spindle tree, etc.) or, con- 
trarily, consume only bark (of spruce in Pechora-Ilych preserve). 
The main form of damage caused to pine by elk in Moscow region is 
the breaking of tops of young pines up to the age of 20 years (59%), 
gnawing of bark (34%), and consumption of shoots (7%) (Dinesman, 
1957). 
267 Under conditions of food scarcity, especially in the latter half of 
winter, elk consume large quantities of arboreal lichens {Bry- 
opogon, Usnea, etc.). In an elk killed on November 19 at Sikhote-Alin, 
arboreal lichens constituted over 50% of the stomach contents 
(Kaplanov, 1948). Sometimes elk rake the snow with their muzzle 
(not more than 20 cm deep) to gather tiny berry shrubs, shoots (most 
often sedge), or even terrestrial lichens. In larch forests in Sikhote- 
Alin consumption of mountain cranberry in one case amounted to 



377 



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378 



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Mean monthly 
temperature 



Fig. 83. Intake of tree and shrub bark by elk depending on atmospheric temperature. 
Lapland sanctuary (from Semenov-Tyan-Shanskii, 1948). 

Note: Roman numerals indicate calendar months. 



268 



31.5% (Kaplanov, 1948). By and large, however, digging food up 
from under snow is not typical of elk. 

Male elk consume in winter up to 9 kg of food (air-dried), female 
elk about б kg, and juveniles up to 5 kg (Semenov-Tyan-Shanskii, 
1948). According to experimental data obtained in Alaska an elk of 
mean annual weight of about 550 kg requires 16 kg of food (air- 
dried) every day (Palmer, 1944). Adult elk in Pechora-Ilych preserve 
in summer consume 30 to 40 kg green fodder (not air-dried) per day, 
in the first half of winter up to 15 to 20 kg, and in March- April, i.e., 
during the "yarding" period, 6 and not more than 8 to 12 kg(Knorre, 
1952 and 1953). 

In winter the water regime of elk, like that of most other ungu- 
lates, is extremely thrifty. The animals lick very little snow since it is 
associated with loss of heat in the organism (Kaplanov, 1948). Feces 
are dry and urination extremely rare, sometimes only once a day 
(Semenov-Tyan-Shanskii, 1948). 

Elk do not visit natural salt licks over much of the range (Euro- 
pean part of the USSR,^^^ western Siberia, Altai, and the Sayans). In 
Irkutsk district visits are rare (Kopylov, 1950), while in Yakutia, 
Amur, and Sikhote-Alin natural salt licks are well-used in the snow- 
free period of the year (Belyk, 1948; Kaplanov, 1948; Abromov, 1949). 
From time to time the animals go to salt licks in winter also 
(Sikhote-Alin). In Sikhote-Alin elk lap turbid water and muddy the 
soil, especially around sedge root tussocks. ^^^ In addition to water- 
In the northern Urals elk in spring and summer go to some definite places in 
marshes where they lap stagnant water with a rusty bloom (Turkin and Satunin, 
1902). 

When mineralized waters rise to the surface around roots of sedges, salts are 
evidently precipitated (Kaplanov, 1948). 



379 

logged solonetzes they also visit marshes (to swallow "liquid peat") 
and dry salt licks where they lap viscous clayey soil and the weather- 
ing crust of tuff; in these environs they eat sphagnum moss 
(Kaplanov, 1948). On marine coasts of the Primor'e territory elk 
consume laminaria and other algae which are rich in salt, lick salt- 
coated stones, and lap sea water (Abramov, 1954). In several regions 
of the USSR elk visit artificial salt licks. In winter in eastern Siberia 
and other regions of the USSR elk avidly lap water rich in organic 
compounds brought to the surface at places by cryoturbation. 

A mineralogical analysis of the composition of salt licks has not 
provided a satisfactory explanation as to why elk visit them (Peter- 
son, 1955; Yu.A. Liverovskii). 

In captivity elk eat cabbage, root crops, potatoes, and other culti- 
vated plants. 

Home range. Except during the period of rut, when males with- 
out females wander extensively, and the period of autumn-winter 
and spring migrations, which do not occur everywhere, elk lead a 
fairly settled way of life, remaining long within a given locality. In 
summer the individual home range of an elk is more extensive than 
in winter, but if the animal has occupied from spring onward a 
home range close to water sources, rich in food, and free from 
danger, it spends the entire summer there; migrations in such cases 
are minimal (Kaplanov, 1935). When the population density is high, 
boundaries of individual home ranges overlap because elk possess 
no "defended territory" from which other animals are driven away. 
The only obvious exception is the female, which during the first few 
days after calving will allow no other elk nearby. 

In Sikhote-Alin preserve in summer elk move several kilometers 
to salt licks but do not travel daily. As many as 28 elk have gathered 
at a single salt lick at one time (Kaplanov, 1948). When salt licks are 
absent such extensive movements of elk outside the limits of their 
permanent habitation are atypical. In the period of rut elk live in 
pairs in a given area (Kaplanov, 1935; Semenov-Tyan-Shanskii, 
1948); their daily movement in general is 5 to 6 km. Single males 
sometimes wander several tens of kilometers daily in search of a 
female (Knorre, 1953). 

In winter, as the snow height increases, daily wanderings of elk 
decrease concomitantly. In March, when the snow cover is 65 to 70 
cm high, daily wanderings are limited to 700 to 900 m. In Lapland 
preserve in early March an adult male elk in a pine forest did not 
move from its 10-hectare home range for five days; a large number of 
animals settled for an even longer period (Semenov-Tyan-Shanskii, 
1948). In the Dem'yanka basin a group of three or four elk in the 



380 

latter half of winter daily traversed 1 to 3 km or sometimes even 
more; only frightened elk move tens of kilometers or more 
(Kaplanov, 1935). In the Pechora basin elk are equally static in win- 
ter. Herds remain for days in an area of a few hectares when the snow 
cover is high. Individual yards there are quite dispersed (Teplov and 
Teplov, 1947). 

In Darwin preserve, when the snow height is 40 cm at the end of 
January, females and fawns move daily about 500 m in a pine-aspen 
forest outside a territory of about 2.5 hectares (Nasimovich, 1955). In 
Voronezh preserve the daily wandering of a male elk which fed in 
269 early December on river banks was 800 m (Nikul'tsev and Predte- 
chenskii, 1957). Similar data on the low mobility of elk in winter, 
especially in the period of "yarding" during deep snow, have been 
recorded for Leningrad district (Sablinskii, 1914), Buzuluk pine 
forest (Knorre, 1939), and several other areas. ^^^ 

In summer elk are widely distributed but gather into herds in 
winter at a few places where food is more abundant and snow less. In 
Lapland preserve in a forest area of 1,000 hectares the average den- 
sity of elk in 1940 was 0.8 and in 1941, 1.0. In yards, however, the 
corresponding values for 1,000 hectares were 3.7 and 4.6, i.e., almost 
five times greater (Semenov-Tyan-Shanskii, 1948). In Mordov pre- 
serve in the mid-1 940's, in spite of a high overall density of ungu- 
lates, elk occupied in winter only 39% of the territory of the preserve 
(Kozlov, 1947). 

Indices of elk population density in different parts of the Soviet 
Union have already been given (see the section "Population"). In 
the latter half of the 1940's these varied on the average from 0.3 to 1 .0 
per 1,000 hectares (Danilov, 1949). At places where elk are protected 
and food still available, population density may be much higher — 
up to 20 to 25 animals per 1,000 hectares (Oka preserve, 1945^^"* and 
Priok-Terras preserve, 1952 to 1955). Experience has shown, how- 
ever, that under these conditions, forests are greatly damaged, winter 
food soon exhausted, and the elk population in subsequent years 
smaller (see "Population Dynamics"). 

Daily activity and behavior. In summer elk are active during the 
day, mainly in the first two or three weeks of June. With the appear- 

^^^According to Semenov-Tyan-Shanskii (1957) the average daily wandering of elk 
in Arkhangel'sk district with a snow thickness of 80 to 120 cm was 3.5 km, going up 
to even 3.9 km for some individuals in the herd. Obviously contradictory to this is 
mention of the habitation of individuals in an area 1.9 km x 0.8 km (152 hectares). 
Some errors have evidently crept into the author's data. 

"''■'in some sections of Oka preserve the population density of elk has reached 34.5 
per 1,000 hectares (L.P. Borodin). 



381 








7r>? 



Fig. 84. Female elk in resting place in water. Position of head with nostrils touching 

the water to avoid possibility of attack by gadflies is characteristic of elk. Elk farm of 

Pechora-Ilych preserve. July, 1959. Photograph by E.P. Knorre. 

ance of a large number of gadflies and horseflies and high daytime 
temperatures, elk take to a nocturnal mode of life. During the day 
they lie up in quiet, cool places where the wind blows strongly and 
blood-sucking flies are few. In montane regions they often ascend 
slopes (Sikhote-Alin, Altai, and southern Urals), emerge into open 
270 zones and large clearings, and move above tree line. Quite often, for 
example in close proximity to populated areas, elk remain hidden 
during the day in thickets of young coniferous plants, in swampy 
alder thickets, and among shrub thickets. At places of little distur- 
bance they remain during the day in open swamps, along shores of 
lakes, and in shoals and river spits, often lying in shallow water and 
at times advancing into water up to their neck. In the warm period 
elk like to remain in wet places; when the spot heats up the animal 
shifts to a new place. 

In the period of mass flights of horseflies elk in Pechora-Ilych 
preserve remain in their beds for about 60% of the day versus slightly 
more than 50% in the period when horseflies are absent (I.S. Turov, 
1953). However, even at places where horseflies are negligible 



382 

(Lapland preserve) elk remain lying for long periods in summer 
(Semenov-Tyan-Shanskii, 1948). Evidently the animals are troubled 
during the day not only by horseflies but also by high air tempera- 
ture, w^hich confirms observations made on captive elk (Knorre, 
1953). 

In summer elk emerge for feeding when twilight sets in and the 
heat has diminished, returning to their beds not later than 6:00 or 
7:00 a.m. They feed in burned-over sections, in bank-side shrubs, 
marshes, and often in water bodies in which they sometimes spend 
much of the day. Elk like old river channels with stagnant pools 
thickly overgrown with dense aquatic vegetation, small streams, and 
shallow bays of lakes. The animals get at aquatic vegetation by 
submerging their head under water (for 20 to 60 sec; Denniston, 
1956). In shallow places they prefer to feed with their ears above 
water. In deep waters they sometimes dive for food (Cherkasov, 1884; 
Petri, 1930; Dmitriev, 1938; and others). Instances are known of 
American moose diving to a depth of 5 m and remaining under 
water for 50 sec or more (usually not more than 30 sec) so that even 
the ripples on the surface had vanished. Feeding on aquatic vegeta- 
tion continues uninterruptedly for 30 to 60 min, followed by a pro- 
longed interval during which the animals stand in water or even 
leave it. Sometimes they recommence feeding within 10 to 15 min, 
however (Peterson, 1955). 

Elk visit salt licks at night and, if not threatened, in the evening 
or morning as well. They visit nearby salt licks daily, as many as 
seven or eight times during the night (Peterson, 1955). They do not 
go to distant salt licks every day but remain close to them, often for 
several days (Sikhote-Alin). In a salt lick elk suck water and semi- 
fluid mud through their lips for 10 to 15 min or even an hour almost 
uninterruptedly. They subsequently feed or lap clean water 
(Kaplanov, 1948). 

Elk easily swim for a few kilometers. They periodically visit 
islands in lakes, remaining 2 to 3 km away from the shore. In 
Lapland preserve 12% of the summer occurrences of elk were ani- 
mals swimming through the lake (Semenov-Tyan-Shanskii, 1948). 
The rate of quiet swimming of an elk is about 2 km an hour ^^^ 
(Kaplanov, 1935). Instances are known of elk swimming across 
Rybinsk reservoir, a distance of 20 km. Similar distances have been 
reported for Scandinavian and American moose (Peterson, 1955).^^^ 

^"Reindeer during migrations swim through rivers at a speed of about 5 km an 
hour. 

^**References occur in literature to instances of elk swimming across the Gulf of 
Bothnia (Kalnin'sh, 1950). 



383 




271 Fig. 85. Swimming elk. Deep lake, Moscow district. November 7, 1958. Photograph 

by V.D. Lebedeva. 

Elk cross a marsh by the shortest route, selecting places where 
land masses rise sharply or where "islets" or ridges occur (Kaplanov, 
1935). Buturlin (1934) has emphasized the surprising ability of elk to 
move through marshes; in swampy places the animal crawls on its 
belly with its forelegs stretched out in front. Tarasov (1956) thinks 
elk are better equipped to negotiate marshes than reindeer. In the 
Altai elk cross marshes along definite tracks, stamping out deep 
trenches 50 to 70 cm in depth in drier parts (V.G. Heptner). 
271 Elk in general are quite sluggish. They avoid running without 
purpose and walk with a long stride, covering without difficulty 
1.5 km in 10 min. Elk running alongside automobiles can gallop for 
a few hundred meters at a speed of up to 35 or even 55 km an hour 
(Peterson, 1955). The speed of an elk in a fast trot without special 
haste is 15 to 16 km an hour (Buturlin, 1934). Elk jump less well 
than other ungulates, clearing a fence 2 m high by thrusting their 
forelegs upward and tumbling heavily on the other side, usually 
breaking the fence in the process. 

To get at food on the ground elk usually have to spread their 
forelegs widely or even go down on their knees. While eating fungi, 
sorrel, red bilberry, and lily-of-the-valley young elk often crawl on 
their knees (Shaposhnikov, 1951; Skuncke, 1949). Adult elk particu- 
larly enjoy browsing the tops of relatively tall herbaceous plants. 

In the period of rut elk are active throughout the day. In winter 
they lie down several times and feed several times as well. Early in 
winter adults lie down four or five times a day and at the end of this 



384 

season, when snow is heavier, eight or ten times. Young animals He 
down somewhat more often than aduks (Semenov-Tyan-Shanskii, 
1948). In the Dem'yanka River basin the periods of rest and feeding 
in winter alternate five to six times in a day (Kaplanov, 1935). 

In Komi Autonomous Soviet Socialist Republic during below- 
freezing weather elk remain in their bed for 75 to 80% of the night 
and 35 to 40% of the day. At a temperature of [minus]* 40 to 50° С elk 
lie down for several hours deeply entrenched in porous snow, above 
which only the withers and head are visible; heat dissipation is thus 
minimized (Knorre, 1953). The opinion that elk are more active 
during cold weather than at any other time (Buturlin, 1934; 
Abramov, 1954) is evidently incorrect. It is highly probable at the 
same time that where persecuted by hunters the rhythm of activity is 
shifted into nocturnal hours. According to Kaplanov (1935) in win- 
ter elk are more active at night than during the day. In strong winds 
and snow-storms elk rest anywhere under a shelter and gather in 
thickets of young coniferous plants. 

At the time of feeding elk spend more time motionless, plucking 
branches and needles, than they do walking; no more than 2 hr a 
day are spent in moving through snow 60 to 65 cm deep. In Lapland 
preserve, when the snow cover is 50 to 90 cm deep, elk feed on 
branches and needles of a single tree or bush roughly five times more 
intensely than at the beginning of winter when the snow is no more 
than 10 to 12 cm deep (Semenov-Tyan-Shanskii, 1948). While feed- 
272 ing elk quite often break up small trees by seizing the top in their 
mouth and bending the trunk by moving forward. The track of the 
animal is invariably away from the tree (Semenov-Tyan-Shanskii, 
1948). Yet many authors state that the animal breaks the tree by 
throwing its chest heavily against it and passing the trunk between 
its legs. 

The weight load on the hooves of elk is considerable while walk- 
ing and, depending on the method of calculation, ranges from 322 to 
749 g per cm^ (Table 2) (E.P. Knorre; Pechora-Ilych preserve). The 
weight load is reduced to one-half when the animal rests, standing 
on its hooves, or while moving over snow and in marshy soil. Never- 
theless elk do get stuck deep in snow, almost to the ground or at least 
to two-thirds of the snow thickness. However, elk movement on 
snow is greatly facilitated by its massive build and long legs. 



*Omitted in Russian text — Sci. Ed. 



385 

272 TABLE 2. WEIGHT LOAD ON TRACK OF ADULT PECHORA ELK (IN g PER 

cm' OF HOOF SURFACE) 







With no allowance 


With allowance 


Sex 


Month 


for additional 


for additional 






support area 


support area 


Male 


October 


749 


434 


Male 


March 


624 


270 


Female 


July 


762 


322 



In the Kola Peninsula adult elk negotiate without difficulty a 
porous snow cover 40 to 50 cm deep. A snow cover of 60 to 70 cm 
usually compels elk to move on old tracks, but if the snow is porous 
the animals can run in it without particular difficulty. Under such 
snow conditions young elk walk in the tracks made by adults. When 
the density of snow cover is 0.20 to 0.22 and the height 85 to 90 cm, 
running adult elk push the snow with their belly and move with 
great difficulty (Nasimovich, 1955). A snow cover of 90 to 100 cm is 
273 critical for elk; in such snow, especially with a frozen crust, winter- 
ing conditions become somewhat difficult. Yet even then, if seldom 
threatened by hunters, elk winter well. In regions where hunting is 
more intense the animals prefer places where the snow cover is no 
more than 70 to 80 cm. 

Elk become aware of an approaching enemy mostly through the 
senses of smell and sound. Vision is relatively poor in this animal 
and its ability to identify an immobile man far away poorer than 
that of a reindeer. Before lying down elk usually turn leeward, make 
a semicircle or loop, and lie anywhere on an elevated site, sometimes 
behind a tree or with their head facing upwind. The approach of an 
enemy in such a position is noticed well in advance (Petri, 1930; 
Kaplanov, 1935; Semenov-Tyan-Shanskii, 1948; Kalnin'sh, 1950; 
and others). In the bed elk sleep or ruminate; they never sleep while 
standing. 

Elk rarely attack man; usually only males in the period of rut or 
injured animals do so. An injured animal is extremely dangerous 
since it can easily kill a man by kicking with its forelegs (Buturlin, 
1934). 

Depending on activity, the body temperature of elk varies widely. 
Under great physical stress it may rise to 41 °C; in winter it is usually 
35.8 to 37.0°C, being somewhat lower at 34.0°C in young animals 
(Knorre, 1953). 

Even moderately large groups of 12 to 18 elk are rare and such 
herds usually break up soon. In summer adult females move with 




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387 

calves, often including those of the previous year, while males and 
barren females live singly and rarely join together in mixed or single 
sex pairs and groups of up to three or four animals. At the end of rut 
some continue to live in pairs, and calves and often 1.5-year-olds join 
them. Sometimes even foreign adult elk join the group. Such a 
group may number from five to eight or even ten elk. 

Some old males after rut live singly throughout winter while 
much younger ones band together in small groups to which barren 
females or 1.5-year-old males sometimes attach themselves. The lat- 
ter, separated from their mothers, are often encountered in pairs. 
The herd instinct is more intense in winter than in summer, and in 
winters of heavy snowfalls than in winters of moderate snows. In 
Pechora-Ilych preserve the herding index from October through 
February (1934 to 1938) varied from 1.7 to 2.7 animals; in March, 
when the snow cover was highest, it rose to 3.7 (Teplov andTeplov, 
1947). Herds break up in spring. Gregariousness is more pro- 
nounced in those parts of the range where population density is 
high. 

Seasonal migrations and transgressions. Elk are significantly 
better adapted to living under conditions of a snowy winter than 
many other ungulates. They often winter in places abandoned by 
roe deer and even red deer. In regions with a mean maximum snow 
cover not more than 30 to 50 cm deep and a short period of heavy 
snow (central region of the European part of the USSR, much of 
Yakutia, and elsewhere), most elk winter in the same locality where 
they occur in summer, performing only minor transitions from one 
area to another. Elk lead a relatively settled life in the Baltic region 
and Belorussia even though the snow there is more (mean maximum 
snow cover not less than 50 cm deep; Rikhter, 1948). In Belovezh 
Forest elk wanderings were recorded in the past (Kartsov, 1903), 
which ceased after the population of ungulates in the forest 
decreased and the Pinsk marshes dried up. 

The behavior of elk differs in places where the mean maximum 
snow cover in winter reaches 70 cm or more and the heavy snow 
period is fairly prolonged. In such localities elk concentrate in win- 
ter in the less snowy sections or move into other regions. 

Seasonal migrations of elk in the northern parts of their range in 
regions of heavy snowfalls are quite extensive. Elk live here in taiga 
plains and wander in winter into less snowy regions, which are more 
convenient for wintering from the viewpoint of habitat (see section 
274 "Habitat"), abundance of tree food, etc. (Arkhangel'sk district, 
Komi Autonomous Soviet Socialist Republic, and western Siberia). 
The length of migratory routes of elk during seasonal migrations in 



388 

these regions sometimes reaches 100 to 300 km (Kaplanov, 1935; 
Knorre, 1953; Nasimovich, 1955; Semenov, 1956; Shubin and Yazan, 
1958; and others). 

Mass seasonal migrations of elk are also distinct in some mon- 
tane regions with a high snow cover where some mountains play a 
role in climatic boundaries. Elk migrate in winter into regions pro- 
tected from snow by "mountain screens". Such types of migrations 
of elk have long been known in the Urals where snow is heavy on 
the western side and relatively lighter in regions east of the range 
divide. The animals move in autumn and early winter from west to 
east, from western Ural to Trans-Ural, and vice versa in spring. The 
southernmost point for which information is available about the 
existence of regular elk wanderings through the Urals is the south- 
ern extremity of the Ural-Tau mountain range (about 53° N lat.), 
while the northernmost point is the upper course of the Ilych (north 
of 63°). The distance between them exceeds 900 km. The territory 
from which elk migrate in winter through the mountain range to 
the east extends in the west from the watershed divide for roughly 
100 km. However, the distance traversed by wandering elk, reckoned 
from end to end, reaches 200 km in some regions of the Urals. The 
density of migrations can be judged from the fact that in the mid- 
1940's from 4,000 to 6,000 elk migrated annually from the Perm 
region alone to winter "beyond the Urals" (Kuklin, 1946). 

In several montane regions with a high snow cover elk move in 
winter into other sections (montane regions of the Kola Peninsula, 
Stanovoy range) or descend along the mountain slopes (the Altai, 
Sayans, and Barguzinsk ranges), wandering right down to the foot- 
hills in particularly snowy winters. The distance covered by elk 
wanderings may add up here to several scores of kilometers. 

In montane regions in the southern Far East (Burein range, 
Sikhote-Alin, and others), during the first half of winter elk occur in 
large numbers at a higher level along mountain slopes than in 
summer, and descend only in February or March immediately after 
the snow becomes heavy (Vetlitsyn, 1894; Emel'yanov, 1927; 
Kaplanov, 1948; Abramov, 1949). The migration of elk in winter to 
higher altitudes in mountains is explained not so much by ecologi- 
cal factors as by the fact that hunting commences at that time of the 
year in the lower regions of the taiga. The presence of large numbers 
of men and dogs results in the migration of elk into deserted regions 
by climbing mountains where, in spite of some snow (40 to 60 cm) 
but not enough to impede their movement, the environs are quiet 
and safe (K.G. Abramov). Other reasons such as the absence in upper 
montane zones of wolves and competing species of ungulates could 



389 

also be incentives for migrations (Nasimovich, 1955). 

In many regions, in addition to wandering elk, some resident 
animals are seen. According to Krylov (1926), in the 1870's elk in the 
region of the Kolva River ( Vishera basin), which had not crossed the 
Urals before mid-December, could not negotiate the range later 
because of high snow. Sighting such a helpless elk in February when 
the snow cover was high, often up to 140 cm, hunters bagged it 
without difficulty. Thus hunting acquired a selective importance 
and helped eliminate from the population animals inclined toward 
a more settled way of life (Nasimovich, 1955). 

The seasonal wanderings of elk, like those of other ungulates, 
depend largely on population density; a small population can live 
year-round on local foods, which would prove inadequate for a 
larger group. During the 1910's in the upper reaches of the Un'ya 
and Shizhim (upper part of the Pechora basin), mass seasonal 
migrations of elk were recorded (Belousov, 1915), which ceased with 
a reduction in population. In 1934 the extensive (1,200,000 hectares) 
Pechora-Ilych preserve was organized, after which the elk popula- 
tion was gradually restored. Yet in the first few years of existence of 
275 the preserve extensive migrations of elk never occurred. Only local 
migrations were observed, with elk congregating en masse in winter 
in river valleys and adjacent watershed areas covered by boreal 
forests, pine groves, and burned-over forest sections (Teplov and 
Teplov, 1947). 

Mass seasonal migrations of elk from the Pechora-Ilych into 
other regions occurred for the first time in 1945 and have continued 
regularly since. Now (1958), part of the elk population winters in 
the preserve and part (evidently a minority) from regions adjoining 
the Ural mountains migrate eastward in winter through the moun- 
tain range. The majority of elk from the main territory of the pre- 
serve migrate in autumn southwest and southeast in the direction of 
the divides of the Pechora,^ Kama, and Vychegda, covering a few 
hundred kilometers. 

The following figures provide a good idea of the magnitude of 
elk migrations in the Pechora basin: in the latter half of November, 
1947, about 600 migrations of elk southwest for a distance of 150 km 
were recorded through the Ilych River, serving at that time as the 
northern boundary of the preserve. In December, through the south- 
ern and western boundaries of the preserve over a stretch of 200 km, 
250 crossings of elk occurred beyond the limits of the preserve in a 
southwesterly direction. Within the preserve itself in that year no 
less than 600 animals wintered, representing new arrivals and local 
elk (Knorre, 1953). In autumn of 1955, 9 km from the farmhouse 



390 

office of the preserve at Yaksha, no less than 1,000 elk moved 
through a section 6 km long (Shubin and Yazan, 1958). 

The transit of elk toward the wintering site is usually gradual 
and continues for two months or more from October (sometimes, 
rarely, even from September) until early December, sometimes from 
November through December or even January inclusive. The com- 
mencement of mass migrations of elk to wintering sites often coin- 
cides with the first significant snowfalls, after which a permanent 
snow cover forms. At other times the initiation of migration coin- 
cides with the period when the snow cover on the summer range 
reaches 25 to 45 cm. Sometimes elk move out for wintering even 
before snows begin, immediately after the first significant cold 
snaps. Other elk remain in summer areas until the snow cover 
reaches 60 to 70 cm and the animals begin to experience considerable 
difficulty in moving around. In most cases they go through the 
period of rut in the same region in which they spent the summer. 

In autumn elk move singly, in pairs, or in small herds. They 
cover 10 to 15 km in a day, sometimes up to 30 km, and then remain 
for some days at a particular place (Dem'yanka basin; Kaplanov, 
1935). In the Pechora basin the first to go are females with calves and 
yearlings, while adult males and females without calves mainly 
move out at the end of the migratory period (Shubin and Yazan, 
1958). 

Spring migrations occur over a very short period, mainly when 
snow thawing is maximum. From the Dem'yanka basin elk leave in 
spring in April; in years with frozen snow crust they move singly 
and in pairs after thaws commence (Kaplanov, 1935). In spring the 
first to leave are usually adult males and females without calves, and 
the last females are with calves (Malaya Sosva and the Pechora 
basin). In case movements have ceased during crusted snow forma- 
tion, then migrations are particularly intense after the crust has 
cleared. In the spring of 1942, from April 29 through May 3, in an 80 
km stretch between Nyagyn and Khangokurt, V.V. Raevskii counted 
200 tracks of migratory elk moving north (Konda-Sosva preserve). In 
some regions part of the elk returned to their summer habitats only 
in June (Nasimovich, 1955). 

Often the routes of elk migrations are extremely regular. The 
animals prefer to move along river valleys; in mountainous country 
they often select slightly longer routes, but with a less rugged topo- 
graphy. In the 1870's elk wandering in winter from the Kova River 

^^'According to Abramov (1954) elk migrate during the course of rut from the 
upper courses of the Bikin. 



391 

beyond [east of] the Urals initially moved southeastward. At Bere- 
zovsk Kamen' their course turned steeply southward parallel to the 
276 mountain range and then northward. Near Chuvamsk Kamen' they 
crossed the Vishera and again moved south parallel to the mountain 
range up to the valley of the Vels River, from whence they turned 
east across the divide (Krylov, 1926). 

Long transgressions of elk, 100 to 200 km or more, into tundra 
and steppe have been recorded repeatedly. Usually they occur in the 
warm season of the year. With the approach of winter, the animals 
move away into forest regions or congregate along river valleys 
where woody vegetation is available. Since the early 1950's elk trans- 
gressions into open tundra around Nenetz have been a regular phe- 
nomenon. The animals arrive there in spring from the south (in the 
spring of 1957 about 20 groups of elk moving northward were 
counted on the Malozemelya tundra), and move in a reverse direc- 
tion in autumn (Skrobov, 1958). Similar migrations of elk (in spring 
into tundra and in autumn into forest-tundra toward river valleys 
where there are genuine forests) have been observed from the latter 
half of the 1950's in the Taza Peninsula (Gromov and Rapoport, 
1958). 

During the last ten years elk transgressions into large towns have 
been reported, probably the result of high population density. 

The colonization of elk in regions from where they had disap- 
peared earlier for some reason or the other is discussed later (see 
"Enemies, Diseases, . . ."). 

Reproduction. Elk generally move in pairs during the period of 
rut and hence many researchers consider them monogamous ani- 
mals (Buturlin, 1934; Kaplanov, 1935; Kuklin, 1946; Belyk, 1948; Kal- 
nin'sh, 1950; Likhachev, 1955; and others). Monogamy of elk is not 
established, however. According to observations made in Pechora- 
ilych preserve, rut in elk (1952 to 1953) lasts for only a few days (two 
to five, more often four or five) (E.P. Knorre).^^* As soon as one female 
exhibits disinterest in mating, the male often looks for another 
which has come into estrus later, thus covering quite a few females 
during the period of rut (Petri, 1930; Knorre, 1949; Peterson, 1955). 

The presence of more males, often two and sometimes three, four 
or even six, near a single female in the period of heat has been 
observed time and again (Naryshkin, 1900; Petri, 1930; Kaplanov, 
1935; V.V. Raevskii). In the Buzuluk pine forest and Pechora-Ilych 
preserve roughly 50% of the males in rut move with a single female; 

^'^Swedish zoologists also point out that rut in elk continues for just a few days 
(Peterson, 1955). 



392 

in 30% of the cases two males move with the same female, in 10% 
more than two males, and in another 10% a lone male with several 
females (two to four) (Knorre, 1949). In Pechora-Ilych preserve one 
male in rut mated with seven females and all were impregnated. 
According to Knorre (1953) elk are polygamous. However, taking 
into consideration the fact that the tendency to polygamy is seen in 
this species only under certain conditions, such as high population 
density and marked predominance of females (because of selective 
shooting of males) or captive conditions, it would be more correct to 
regard elk as facultatively polygamous animals. 

Combats are quite common among males over females, ending 
sometimes in the death of one or the other contestant (Teplov, 1948; 
Knorre, 1953; Nikul'tsev and Predtechenskii, 1957). Elk with body 
wounds and scars in the period of rut or immediately thereafter are 
quite common. Instances are known of several males attacking a 
bull that is accompanying a female (Kaplanov, 1935). 

Some elk form pairs even before the commencement of rut — in 
the latter half of August. The male, moving usually behind the 
female, emits a sound generally described as a "moan," until the 
female is sexually aroused. This "moan" resembles more a hollow 
bellow, is far less audible than the call of a deer, and is usually heard 
for not more than 0.5 to 1.0 km, but over much longer distances 
under favorable conditions. Males "moan" at dawn and evening, 
rarely at night, and even more rarely during the day. 

A sexually aroused male breaks the branches and tops of small 
trees with its antlers (such damaged trees can be seen even before the 
beginning of rut when elk clean their antlers), sometimes digs pits 
277 with its hooves, and eats soil wetted with urine of females. Males 
exude a characteristic odor. Females, but more particularly males, 
abandon their usual caution in the period of estrus. Males become 
aggressive and permit the easy approach of man. The daily cycle of 
activity is disrupted. In males the hair becomes disheveled, the eyes 
bloodshot, and the neck thickens (Kaplanov, 1935). Males eat little at 
this time and lose up to 17% of their weight compared to that of the 
period before rut (Knorre, 1956). More aggressive males in the period 
of rut drive calves away from females and move singly, joining the 
female later. Yet in several regions during the period of rut calves 
and females with a single male are not an uncommon sight (Petri, 
1930; Kaplanov, 1935; and Semenov-Tyan-Shanskii, 1948). 

A male will mate with a receptive female several times in the 
course of a day. Mating is very brief, lasting only a few seconds 
(Knorre, 1953; Peterson, 1955). 

The entire period of rut of elk, from moaning to the last mating, 



393 

extends over 1.5 to 2.0 or more months. Mating is limited to much 
shorter periods — a month or slightly more (the majority of elk usu- 
ally mate for 10 to 20 days) but some (usually younger individuals or 
those injured, etc.) mate much later. In Lapland preserve a newborn 
elk was found on July 4. The gestation period of elk is 225 to 237 days 
(Buzuluk Bor and Pechora-Ilych preserve; Knorre, 1953) and in some 
cases not less than 240 days (Perm zoo, Podososov, 1935; Serpukhov 
[Game]* Farm, Semenov-Tyan-Shanskii, 1948). Thus the female which 
calved so late had mated between November 5 and 20. In Quebec 
(Canada) a newborn calf was found on August 14, which means 
mating occurred not earlier than mid-December (Moisan, 1956). 

In the southern and central parts of the range rut in elk is much 
shorter in duration than in the north. After a severe and snowy 
winter and an unfavorable summer (drought, etc.), rutting commen- 
ces more gradually than in normal years (Abramov, 1949 and 1954). 
The duration of rut is undoubtedly influenced by the nature of 
autumn weather but authors do not agree on this fact. Rut in elk 
commences earlier and also usually terminates earlier in the south- 
ern and at some places (mainly within European USSR) in the 
central parts of the range. There elk mating, at least in some years, 
occurs even in the first ten days of September (southern Belorussia, 
Moscow and Saratov districts, Buzuluk Bor, Mordov preserve, 
Rybinsk reservoir, and Sikhote-Alin), and in some cases even at the 
end of August (Saratov district). Rut in elk in these regions termi- 
nates quite often at the end of September or early October and more 
rarely on October 10 to 15. 

In the northern parts of the range and also in regions with rela- 
tively severe climate, prolonged winter, and late spring, mating usu- 
ally commences not before the second decade in September, mass 
mating occurring from September 15 to 20 through October 5 to 10 
(northern part of Scandinavian Peninsula, Karelia, Lapland, 
Pechora-Ilych and Konda-Sosva preserves, Dem'yanka River, Yaku- 
tia, northern Amur region, etc.). Rut in elk in these regions termi- 
nates by October 15 to 25 but some animals mate sometimes even 
later. Much later periods of rut in regions with cold climate 

*Word omitted in Russian text — Sci. Ed. 

^''''Мапу references to the period of rut in elk conflict with data by the same 
researcher on the time and birth of calves. The commencement of the period of rut is 
usually reckoned from the time of pairing, most often from the first instances of male 
moaning, and more rarely the first actual mating. Furthermore, these are not pre- 
cisely recorded. Hence data on the period of estrus in elk should be handled with more 
caution and corrected as far as possible to accord with dates of the appearance of 
young. 



394 

should be regarded as arising due to natural selection; with early rut 
and calving, mortality among the young if cold spells return or 
predators attack, especially bears (see "Enemies, Diseases, . . .") will 
be much higher than among those born at a much later period. 
278 It is highly probable that the prolonged season of rut in some 
parts of the range, and hence of calving, is also of adaptive value and 
caused by fluctuating climatic conditions in spring. With respect to 
the Kola Peninsula, where the period of rut in elk is highly pro- 
longed and the periods of snow cover vary greatly, the above conclu- 
sion becomes self-explanatory. Periods of rut vary widely in different 
years. In Pechora-Ilych preserve mating of elk in 1952 occurred from 
September 23 through October 10 and in 1953 from September 12 
through October 12 (E.P. Knorre). 

With the exception of some individuals, probably those from late 
calving or those which are sickly and lag behind in growth, all elk 
are generally capable of mating in their second autumn. However, 
under natural conditions many females evidently mate only in the 
third autumn. Mating by males before their third or fourth year is 
quite rare since they cannot compete with older bulls. Most younger 
animals become sexually aroused at a much later period than older 
ones. In 1933 older elk in Dem'yanka River basin became aroused 
from September 22 through October 12, while younger elk displayed 
excitation from October 4 through 20 (Kaplanov, 1935). 

At places where elk are hunted often calving usually takes place 
in a less accessible or dense and less frequented spot — dense thickets, 
in young plantations, sometimes along river banks, in marshes, 
islands, peninsulas, etc. In Lapland preserve elk calve everywhere: in 
marshes, alongside creeks, in forests, in burned-over sections, and 
even in montane tundra (up to 500 m). If snow is considerable in the 
period of calving, calves are born in thawed patches (Semenov- 
Tyan-Shanskii, 1948). Before calving and in the first few days after 
it, the female usually drives away yearlings which have remained 
close to their mother throughout that period (Lenberg: cited from 
Peterson, 1955). 

Calving is earliest in the southern parts of the range and at some 
places also in the central region (European part of the USSR) — from 
the end of April to the commencement of May and through May 20 
to 25 (more rarely right up to early June). In some years some 
females calve there from April 20 or even at the beginning of that 
month. In Moscow and Vladimir districts the majority of elk calve 
around May 1 (V.G. Heptner). In the Rybinsk reservoir region (Dar- 
win preserve) calving of elk occurs from around May 10 but new- 
borns were seen in 1951 even by April 20 (Kaletskaya, 1953). In the 



395 

Saratov district the first newborns were seen in 1946 on April 5 and 6 
and in 1958 on April 1 and 4; the last calves were found mid-May 
(Martynov, 1959). 

Calving of elk occurs in Pechora-Ilych preserve, Karelia, Yaku- 
tia, and western Siberia from around May 10 (in rare cases, earlier) 
through June 10. In Pechora-Ilych preserve in 1952 young were born 
from May 27 through 31 and in 1953 from May 11 through 20 (E.P. 
Knorre). In L.apland and Konda-Sosva preserves, and also in the 
northern parts of the Scandinavian Peninsula, the majority of births 
take place in the last ten days of May and the first half of June. In 
Lapland preserve two instances have been recorded of calving of elk 
at the end of June to early July, which should be regarded as a fairly 
rare phenomenon. At places where elk undergo considerable sea- 
sonal migrations, calves are usually born in summer residences; in 
the event of an extremely snowy and prolonged winter calving often 
occurs in Avintering sites (Komi Autonomous Soviet Socialist 
Republic; G.G. Shubin). 

The female elk usually gives birth to one or two calves. Two 
calves, rarely one, are characteristic of the Baltic region, central 
regions of the European part of the USSR, Novosibirsk district, the 
Pre-Baikal region, left bank of the lower Amur, Okhotsk coast, and 
Yakutia. One or two calves are equally common in Belorussia, Bal- 
tics,* Lapland preserve, Saratov district, Buzuluk Bor, Dem'yanka 
River basin, and evidently also in the central Urals. In an over- 
whelming majority of cases one calf and very rarely two are common 
in Trans-Baikal, Dusse-Alin, and Sikhote-Alin. 

In Lapland preserve 25 females with calves were found from the 
end of May through August inclusive in different years; of these, 44% 
279 had two calves and 56% only one (Semenov-Tyan-Shanskii, 1948). In 
Buzuluk Bor over a five-year period elk with calves were sighted 79 
times: 42% had two calves each and 58% one each (Knorre, 1939). In 
one of the Saratov district forests, of 13 females with calves observed 
in 1947, six had two calves each and the rest one each. In Pechora- 
Ilych preserve from 1938 through 1946, 108 females with calves were 
observed; 47% had two calves each and 53% one each (Teplov, 
1948). Of the 25 pregnant elk caught in Novosibirsk district, 23 
(92%) contained two embryos each and only two one each (Likha- 
chev, 1955). Kapianov (1948) during a three-year study in Sikhote- 
Alin detected 23 females with calves, all of whom had only one calf 



*Inconsistency in Russian text — Sci. Ed. 

''"According to Knorre (1953) elk of Pechora-Ilych preserve produce two calves 
more often than one. 



396 , 

each; two calves were rare exceptions. Only 10 to 25% of American 
moose in different parts of the range produce two calves each and the 
rest one each (Peterson, 1955). In Sweden three embryos were 
detected in a lone elk (Lenberg; cited from Peterson, 1955). A single 
female with three calves is considered a great rarity among American 
moose (Peterson, 1955). 

Differential fertility among elk in different parts of the range is 
quite evident, but the reasons for this phenomenon are far from 
clear, particularly since accurate numerical data are inadequate and 
those which are available are not always comparable. Some 
researchers take into consideration encounters of elk with offspring 
only in the summer months and others throughout the year, which 
is less exact (see below), etc. 

The proportion of single to twin calves largely depends on the 
population composition of elk; moreover, it varies widely in differ- 
ent years. In a given locality fully adult females often produce two 
calves, while younger females deliver only one. In the first few years 
of organization of Buzuluk Bor preserve, young elk predominated in 
the population. As a result, from 1933 to 1936 the percentage of two 
calves with a single female was only 10 to 31%, which rose to 45 to 
57% from 1937 to 1940 (Knorre, 1949). In Pechora-Ilych preserve the 
average "offspring per female in different years ranged from 1 .2 to 
2.0. A greater number of offspring were noticed after normal and 
relatively mild snowy winters and a smaller number after severely 
snowy winters (Teplov, 1948). 

Among twins, one may be a female and the other male (47%), 
both males (30%), or both females (23%). The second calf of the twin 
set often dies. In Buzuluk Bor, according to data over a seven-year 
period, twins in May averaged 57%, in May-June 52%, May-Sep- 
tember 46%, and over a 12-month period only 40% (Knorre, 1949 and 
1953). 

Some elk remain barren. In the Primor'e region barrenness is 
rare (Abramov, 1954), while in other parts of the range it reaches 30 
to 40% (Pechora basin) or more. Barrenness is generally more com- 
mon if elk hunting commences before the conclusion of the period 
of rut (Skuncke, 1949), or in the event of a severe winter when the 
period of rut is prolonged beyond the normal period. Among Ameri- 
can moose usually not more than 50% of animals older than two 
years fawn every year, the rest fawning in alternate years. In New- 
foundland, Ontario province, and Alaska the number of barren 

^" Whether calves in this case died in the first few days after birth or whether 
females with twins aborted more often has not been indicated by the author. 



397 




280 Fig. 87. Newborn calf on the Chuna tundra. Lapland preserve. July 4, 1939. Photo- 
graph by O.I. Semenov-Tyan-Shanskii. 



280 



animals may reach 60 to 65% of the population (Peterson, 1955). 

Growth, development, and molt. Observations on elk born in 
captivity confirm that within the first 10 to 15 minutes after birth 
they can stand on their legs but fall down immediately. The pelage 
and umbilical cord are wet on the first day. By the second day calves 
move better even though their legs are still gangly and sometimes 
uncoordinated. From the third day they move freely and on the fifth 
day it is difficult to catch them; by the tenth day, they are not inferior 
to their mothers in the use of their legs; and at the age of two weeks 
they are excellent swimmers (Knorre, 1953; and others). Under natu- 
ral conditions calves remain in one place for at least a week. When 
the mother goes away to feed or runs away on sighting a man, the 
calves remain hidden in the grass or bushes; the mother makes no 
attempt to protect her young from man (Kaplanov, 1935; Semenov- 
Tyan-Shanskii, 1948; Knorre, 1953). 

Lactation continues for 3.5 to 4.0 months, i.e., roughly until the 
commencement of the period of estrus. Some females, those which 
have not gone into heat, continue to lactate in November- December 
or even later (Kaplanov, 1935; Knorre, 1953; Likhachev, 1955; Peter- 
son, 1955). A barren female, killed in Pechora-Ilych preserve at the 
end of December, yielded 200 g milk. The calf with her weighed 43 



398 

kg more than any other in the preserve (E.P. Knorre). On the elk 
farm of Pechora-Ilych preserve one female yielded 150 to 200 liters 
milk during the period of lactation with a maximum of 2 or even 3 
liters per day (June to early July) ; the daily yield was minimal at 
the commencement and termination of lactation. The fat content of 
milk in May-June was 8 to 10% and even up to 13% (Knorre, 1953; 
G.G. Shubin). Compared with cow's milk, milk of elk has 2.4 times 
more fat and ash, 5.0 times more proteins, but 1.6 times less lactose 
(Krott, 1956). Calves begin to eat green fodder around two weeks of 
age or a few days later (Podsosov, 1935; Shaposhnikov, 1951). In 
captivity calves attempt to nibble green leaves at the age of two or 
three days. Calves isolated from mothers at the age of 1 .5 months and 
later fed on green fodder only, grew quite normally and were not 
inferior in growth to other fawns (E.P. Knorre). 

A study of 56 calves raised in Pechora-Ilych preserve and Buzuluk 
Bor revealed that the weight of newborn female calves varies from 6 
to 14 kg and of males 8 to 16 kg. Calves from paired litters generally 
did not weigh more than 10 kg. Calves weighing 6 to 9 kg were 
usually very weak and quite often died subsequently (Knorre,1949 
and 1956; and others). Data on the weight of newborn calves from 
other parts of the range consist of occasional weighings (Lapland 
preserve, Serpukhov Game Farm, Dem'yanka River basin, and 
281 Novosibirsk and Irkutsk districts) and do not differ from the forego- 
ing. Data are not available on the weight of newborn calves of elk 
from northeastern Siberia where elk are largest in size in the USSR. 
In Scandinavia the normal weight of newborn calves is 10 to 16 kg 
and of twins sometimes 6 kg each (Skuncke, 1949). 

Calves put on weight very rapidly and within six months their 
weight rises roughly tenfold, with an average of 120 to 130 kg, and in 
better developed animals 160 or even 206 kg. Weight increase in 
calves is relatively low in the first 1.0 to 1.5 months, when milk 
forms their food, as compared to the rise in the subsequent two 
months when large amounts of green fodder are consumed. In July 
the mean daily weight increase in Pechora and Buzuluk calves 
approaches 2 kg (Knorre, 1949 and 1956; and others). The mean 
daily weight increase in American calves in their first month is 450 
to 900 g and in the second month 1 ,300 to 2,250 g (Trippensee, 1948). 

From autumn weight increase slows down and by early winter 
when calves switch over completely to browse, it is even more 
retarded (southern parts of the range) or ceases altogether. In 

"^Improved methods of milking elk later raised the yield to 430 liters in a season 
(E.P. Knorre). 



399 




Fig. 88. Elk calf 25 to 30 days old. Southern part of Khabarovsk territory. July, 1957. 
Photograph by G.F. Bromlei. 

Pechora-Ilych preserve the weight of calves remains unchanged 
from early winter right up to the cessation of the yarding period and 
spring m.olt, or even decreases in the case of a snowy and prolonged 
winter. Thus calves at the age of about one year weigh as much as at 
the age of six months and sometimes even less. Only in elk which 
have not undergone estrus and continue to lactate in winter, which 
rarely occurs, do calves, at least at the beginning of winter, put on 
weight even in the north (Knorre, 1956). 

The height at withers of new-born calves is 70 to 90 cm, at the age 
of two months 105 to 110 cm, at four months 125 to 130 cm (Butur- 
lin, 1934), in the winter of their first year 135 cm, and in the second 
year up to 155 cm. Adults stand 160 to 216 cm at the withers, more 
often around 175 cm (Knorre, 1953; and others). On the elk farm of 
Pechora-Ilych preserve calves usually do not grow up in height after 
October right up to spring, the winter-spring growth stabilization 
being even more distinctly manifested than that of weight (Knorre, 
1956). Male calves at the Yakutia experimental station at the age of 
one month measured 107 cm at the withers and females 105 cm; at 
the age of three months, the corresponding heights were 120 and 1 17 
cm, six months 139 and 132 cm, nine months 146 and 145 cm, and 
twelve months (females) 151 cm (Popov, 1939). The growth of these 
calves and their weight increase continued even in winter. The 
weight increase in elk older than a year is given in Table 3. 



400 

182 In their second summer elk continue to put on weight and under 
particularly favorable conditions (cool and rainy summer and few 
blood-sucking flies) attain 150 kg or more, so that their weight by 1 .5 
years of age has often doubled. Some elk weigh as much as 350 kg. 
The relative weight increase in elk is invariably maximum in their 
first year and the absolute weight increase, depending on weather 
conditions in summer, can be maximum in their first or second year. 
The weight increase slows down in the third year and the animal 
attains full physical stature in the fourth year. Later the weight of 
adult elk undergoes fairly regular seasonal changes, reaching 80 kg 
or more in amplitude and constituting up to 20 to 25% of the maxi- 
mum weight of the animal in a given year. The maximum weight of 
elk is recorded at the end of August to early September and the 
minimum at the end of April to early May. During the period of rut 
males lose up to 17% of their former weight and in the subsequent 
winter three to five times less. In female elk weight reduction in the 
cold season of the year occurs very gradually; during the period of 
estrus, by November, they lose not more than 5% of their original 
weight (Knorre, 1956). 

TABLE 3. BODY WEIGHT (kg) OF ELK IN THE EUROPEAN PART OF 
THE USSR AT DIFFERENT AGES (E.P AND E.K. KNORRE, 1956) 



Sex 



New- 


0.5 


1.5 


2.5 


3.5 


4.5 years 


born 


years 


years 


years 


years 


and older 



Male 8-16 66-200 171-278 222-347 275-375 357-619 

Female 6-14 69-140 175-331 227-345 275-360 347-406 



Observations made in Sweden show that after the fourth or fifth 
year female elk do not put on weight, while males do not usually 
attain maximum weight before their tenth year (Skuncke, 1949). 

Within a given age group weight variability is exceptionally 
high, as a result of which animals of different ages often weigh the 
same: males at the age of 1.5 to 3.5 years weighed in around 275 kg, 
while some 1.5-year-old females and animals aged 2.5 and 3.5 years 
weighed up to 300 kg (E.P. Knorre). 

Data on the weight of elk in Siberia and the Far East are frag- 
mentary but conform quite well within the range shown for the 
European part of the range. The maximum known weight of a 
Siberian elk (male) is 655 kg (Yenisey basin) and of a European elk 



401 

619 kg. One male in over a hundred elk killed between 1903 and 1912 
in B. Petersburg province weighed 619 kg; the rest weighed no more 
than 477 kg each (Sablinskii, 1914). The weight of the largest bull in 
Buzuluk Bor was 563 kg and in Pechora-Ilych preserve roughly 500 
kg; adult elk there usually range from 300 to 450 kg (Knorre, 1953 
and 1956). 

At places of intense hunting of elk maximum-sized animals are 
rarely sighted since most are taken in their first few years of life. Of 
over 100 elk killed in winter in southern Karelia not one weighed 
more than 311 kg (Knize, 1935). The maximum weight of a male 
caught in the Dem'yanka River basin was 422 kg and of a female, 
370 kg (Kaplanov, 1935). The usual weight of elk in eastern Siberia 
is 320 to 400 kg and very rarely (males) up to 480 kg (Cherkasov, 
1884). Eleven elk caught in the Pri-Amur region weighed 260 to 
320 kg (Abramov, 1954). Males caught at the end of September in 
283 Sikhote-Alin weighed 400 kg each (Kaplanov, 1948) although Ussuri 
elk are considered the smallest in the USSR. Exact data are not 
available on the weight of the largest-sized elk of the USSR, i.e., 
from northeastern Siberia; the weight of males there in the bloom of 
their strength quite often reaches or even exceeds 600 kg. 

In calves four to five months of age visible lumps have developed 
under the skin on the head by their very first autumn and antlers 
protrude from the end of April to early May through June inclusive, 
i.e., at the end of their first to early in their second year. Soft antlers 
harden only at the end of July or in August; their velvet gradually 
shrivels and is jettisoned by rubbing the antlers against small trees. 
These antlers measure 20 to 28 cm, sometimes up to 32 cm (Buturlin, 
1934), and are often in the form of stubs without tines, bifurcate only 
in exceptionally rare cases. Young animals shed their antlers after 
older ones, usually only in February-March and sometimes even in 
April. The second antlers of elk form at the commencement of their 
third year and are bifurcate. Antlers with well-developed palmate 
portions usually form only in the fifth year. Later, given favorable 
conditions the weight of antlers increases, the palmate portion 
enlarges, and the number of tines increases. The weight of a pair of 
large antlers may reach 15 to 20 kg (Buturlin, 1934) or even more 
according to some authors. 

New antlers commence growth in adult elk in April in the south- 
ern parts of the range and only in May in the north. Antlers attain 
full development at the end of June to the first half of July (quite 

^'^Kopylov (1950) gives the weight of 300 to 400 kg, exceptionally 600 kg, for the 
Irkutsk district 



402 

often in June in the southern parts of the range). Thus their growth 
continues for 2.0 to 2.5 months. As long as the antlers are soft, they 
are very sensitive to impact and to insect bites. Hardening of antlers 
occurs in July; the tips of antlers remain soft most of the time in the 
form of round knobs and become sharp only later. By the end of 
August to early September velvet has been shed but this process 
occurs in Kola Peninsula only from the end of August through 
mid-September. At the commencement of the period of rut the 
antlers of adult elk are invariably free of velvet. In Sikhote-Alin 
young elk with velvet remnants have been sighted as late as Sep- 
tember 17, while the antlers of old animals in some years are clean 
even by August 26 (Kaplanov, 1948). 

Adult elk shed antlers from November (more rarely from the 
latter half of October) through December, sometimes extending up 
to early January.^'"* In the Kola Peninsula and Yakutia elk shed 
antlers mostly in December. Three-year-old elk shed antlers in 
January-February. In old elk the palmate portions shrink, become 
lighter, and often even the number of tines is reduced. Under unfa- 
vorable conditions deterioration of antlers is seen even in animals no 
older than six to eight years (Knorre, 1953). 

Elk are born with well-developed milk incisors and visible pre- 
molars. The formation of permanent incisors in Russian elk is com- 
pleted roughly in their 18th month (Knorre, 1949). In American 
moose the first molar begins to erupt at the age of 10 to 14 weeks 
(that on the lower jaw somewhat earlier than on the upper) and 
functions fully at 4 to 6 months; the second molar begins to erupt at 
6 to 8 months. At 13 to 16 months elk usually lose all the milk 
premolars and formation of molars ceases by 16 to 19 months (Peter- 
son, 1955). 

Young elk have rusty-colored wool, differing sharply from the 
grayish-brown of adult elk; their legs are not lighter in color than 
the trunk. Shedding of juvenile hair occurs from early August 
(slightly later in the north). By the middle or end of September 
young elk acquire the pelage of adult elk; at that time the legs 
become lighter in color and the trunk a dark brown (Knorre, 1953). 
In Lapland preserve elk molt in September, but young ones in juve- 
nile coat have been sighted as a rarity even in November (Semenov- 
Tyan-Shanskii, 1948). 

Data on the molt of adult elk under natural conditions are very 
scanty. One reason for this is the extreme similarity in color of 

^^"Kopylov (1950) has evidently erred in stating that adult elk in the Irkutsk district 
do not shed antlers before February. 



403 

284 summer and winter coats, the former being only slightly darker than 
the winter dress. Elk molt once a year, in spring. By March the 
winter coat is perceptibly worn out and its luster lost. Guard hair 
begins to fall out at the end of March to early April and the underf ur 
in the latter half of April. Molt commences from the head and legs 
and the last to molt is the back. Molt is most intense in May-June; 
however, females which have calved molt in June and the first half 
of July. In the northern parts of the range molt is delayed a week or 
two compared to more southerly parts. The first to molt are males 
and barren females and the last, females which have calved, and 
emaciated and sick animals. In Sikhote-Alin adult males molt early 
in July or even earlier, while females do so only toward August 
(Kaplanov, 1948). Normally fed males and females killed in the 
Dem'yanka River basin from July 16 to 20 had totally molted, while 
a female, still rearing her young and very emaciated, retained the 
remains of winter coat even on July 25 (Kaplanov, 1935). 

The period of molt is difficult for elk, especially for the young. 
At that time body weight falls sharply or weight increase is arrested. 
Some young animals which have come through a severe winter lose 
up to 30 kg in weight during the spring molt (Knorre, 1949). 

From the latter half of July to August elk sport a short lustrous 
summer coat; hair on the abdomen is very sparse. The skin is 
slightly thinner than in winter. In August the underfur begins to 
grow and guard hair elongates. During October,^^^ or slightly ear- 
lier, elk sport a winter coat. 

The strength of elk is at its peak from the age of 6 to 12 years. The 
prevalent view among Soviet zoologists is that elk live for no more 
than 20 years (Buturlin, 1934; Kalnin'sh, 1950; and others). How- 
ever, a male elk tagged in Sweden as a calf and released was caught 
20 years later and found to be in good health with antlers bearing 1 1 
and 12 tines (Lenberg; cited from Peterson, 1955). In the Stockholm 
zoo a female elk calved at the age of 21 years but the calf did not 
survive (Skuncke, 1949). Judging from these data the potential lon- 
gevity of elk is greater than 20 years and possibly exceeds 25 years as 
suggested by Cherkasov (1884). However, an overwhelming majority 
of the animals perish much earlier. In the elk population of 
Lapland preserve those older than 10 years constitute no more than 
3% (Semenov-Tyan-Shanskii, 1948). 

Enemies, diseases, parasites, mortality, competitors, and popula- 
tion dynamics. Among various predators attacking elk the most 

^^'An adult elk caught in the Dem'yanka River basin on September 27 had a dark 
[summer] coat (Kaplanov, 1935). 



404 

important are wolf and brown bear, the former mainly in the least 
snowy parts of the range and the latter in very snowy taiga. 

A healthy adult elk is a difficult quarry for a wolf, the hunt 
posing a great risk to the predator: an elk kicking with the hooves of 
its forelegs can crush the head of the predator. Therefore, unless 
forced to, wolves do not attack elk. When other species of ungulates 
or cattle are available, elk are relatively rarely threatened by wolves. 
In 17 European, Siberian, and Far East preserves, several hundreds 
of remains of wild ungulates torn to pieces by wolves were recorded 
from 1945 to 1949; of these, only 5% were elk even though they were 
numerous (Kozlov, 1953). In Sikhote-Alin preserve, where other var- 
ieties of ungulates are also quite numerous, wolf attacks on elk have 
not been reported (Kaplanov, 1948; Abramov, 1954). Instances are 
known when females with calves in the summer approached within 
100 m of a wolf's lair with litter, or lived from summer to spring in a 
region in which packs of wolves hunted, and suffered no attacks by 
these predators (Martynov, 1951; Peterson, 1955). 

At the same time, when other food is scarce wolves can be very 
dangerous to elk and the resultant population losses sometimes very 
high. In 70 samples of wolf feces collected in Oka preserve, elk 
remains were seen in 77% (Borodin, 1940). In autumn-winter data 
285 on the food of wolves in Darwin preserve (Rybinsk reservoir, 1948 to 
1951; 90 feces), elk remains were found in 78% of samples. At this 
same place and in the same years, 11 young and 10 adult elk car- 
casses torn apart by wolves were found in summer; the correspond- 
ing figures in winter were 9 and 12. Thus, of the total number of elk 
killed by wolves, young animals accounted for almost 50% (Kalets- 
kaya, 1953). Wolves are responsible for considerable losses of elk 
in the Kalinin district (V.N. V'yunkov). 

Young animals, which wolves attack mainly when they stray 
away from their mother, come first in the order of elk decimation; 
juveniles, just when they have taken to independent living, come 
next; and sickly and emaciated adults last (Buturlin, 1934; Murie, 
1944). Wolves usually attack adult elk only in packs and avoid fron- 
tal attack. When wolves which have specialized in hunting elk later 
attack cattle or horses, they hold the prey by the groin and not the 
neck as commonly done (Kalinin district; D.P. Bengard). The dam- 
age to elk by wolves is higher in snowfree and least snowy periods of 
the year. Wolves find it difficult to overcome not only adult elk but 

In the national parks at Jasper and Banff (Canada) elk calves represented 100% of 
prey killed by wolves in summer, and just over 50% in printer (Cowan; cited from 
Peterson, 1955). 



405 

even juveniles in soft and deep snow. 

As a result of intense development of taiga regions in the Soviet 
Union in the last two decades, large forest areas have been felled and 
roads laid. Wolves have penetrated some of these regions of highly 
snowy winters, where formerly they were absent, and the damage 
inflicted on elk by these predators is steadily increasing. Instances 
are known of wolves which previously had never attacked elk later 
mastering the technique of hunting and killing them in large 
numbers (Perm district; Semenov-Tyan-Shanskii, 1948). 

Elk suffer great damage from brown bear mainly in the highly 
snowy taiga regions, especially in spring when the predators emerge 
from their dens, while the snow cover is often deep. Deprived of 
other food sources, bear chase elk on pack ice. The chase sometimes 
covers several kilometers. In Lapland preserve the spring feces and 
stomachs of bear revealed elk remains in 35% of the samples; such 
finds were seven to nine times lower in summer and autumn. Of the 
20 elk killed by bear in the Lapland preserve, 54% were adults, 25 to 
33% animals in the second year of life, and 13 to 21% calves. Among 
the elk killed, one was pregnant with two calves and another in the 
process of calving (Nasimovich and Semenov-Tyan-Shanskii, 1951). 
In Pechora-Ilych preserve 10 deaths of elk caused by bear have been 
established. Of the dead, six were adults, one in the second year, and 
three calves (Teplov, 1953). Deaths of elk caused by bear have been 
recorded in several other regions of the country. According to 
Kaplanov (1935), in the event of an early spring, bears can destroy 
much of the calf crop of elk (Dem'yanka River basin). Evidently a 
high bear population could greatly affect the growth of an elk popu- 
lation. Instances of elk killing or injuring an attacking bear have 
also been described (lokhel'son, 1898; Teplov, 1953; and others). 

Other predators are not significant although some researchers 
hold that wolverine inflict as much or even greater damage on elk as 
bear (Teplov, 1948; Knorre, 1953). Young (up to one or two years) 
and pregnant females and injured or intensely emaciated elk suffer 
most from wolverine (Kozhanchikov, 1924; Teplov, 1948 and 1955). 
Among elk killed in Pechora-Ilych preserve by wolverine, four were 
under one year (one lame), two of the eight adults previously injured 
by poachers, and one chased a long time by a lynx before the wol- 
verine attack. Three pregnant elk chased by wolverine aborted and 
the predator ate the fetuses. Most elk are caught by wolverine in 
February- April by an attack from ambush or by an active chase 
(Teplov, 1955; V.P. Teplov). In Lapland preserve not one instance 
of wolverine attack on elk has been observed; the remains of elk in 
wolverine feces are the result of scavenging (Nasimovich, 1948). 



406 

286 Probably in other parts of the range wolverine is also more often a 
scavenger than a true predator of elk. 

The lynx is usually considered an enemy of elk (Cherkasov, 1884; 
Buturlin, 1934; Kaplanov, 1935; Belyk, 1948; and others) but actual 
data have not confirmed this assumption. In Sikhote-Alin and the 
Altai attacks of elk by lynx are not known (Kaplanov, 1948; Dul'keit, 
1953), but such instances have been recorded in Pechora-Ilych pre- 
serve (Teplov, 1955). Analysis of tiger feces in Sikhote-Alin revealed 
elk remains (Kaplanov, 1948); elk constitute 10% of the victims of 
tiger in this region (G.F. Bromlei; Sludskii, 1953). No reliable 
reports of attack on elk by other predators are known. 

Despite the endurance of elk in deep snow, in very snowy win- 
ters, especially if the animals are persecuted in that period, popula- 
tion decimation rises notably. After the snowy winter of 1894 to 1895 
in the Kolyma basin elk were so severely emaciated by spring that 
they could barely stand on their legs and calves dying from hunger 
were found (lokhel'son, 1898). A considerable number of elk died in 
Siberia during the extraordinarily snowy winter of 1913 to 1914 
(Grum-Grzhimailo, 1914; and others). In Pechora-Ilych preserve, 
where the snow cover in some years reaches 1 10 to 120 cm or more at 
wintering sites, many elk are severely emaciated by spring. In parti- 
cularly snowy winters death due to wolverine attacks increases, abor- 
tions are more frequent, newborns have a low body weight, are very 
weak, and soon die; ultimately the number of young in the popula- 
tion perceptible drops (Teplov and Teplov, 1947; Knorre, 1953 and 
1956; Teplov, 1955; and others). After extremely snowy and pro- 
longed winters, females calve in wintering sites where a large 
number of young evidently perish on their migration to the north 
(G.G. Shubin). 

A prolonged spring with cold spells and snow, like autumn rains 
with frost and wind, is unfavorable for young elk. Population deci- 
mation is high in such years. In summer the most favorable condi- 
tions for the well-being of elk are cold and relatively rainy weather^'^ 
and few blood-sucking flies. When the latter are excessive, some elk 
do not gain weight in that period (Knorre, 1953; and others). 

Elk swim well but instances of their death in water are neverthe- 
less not rare. In Pechora-Ilych preserve a total of 50 deaths of elk over 
a number of years have been recorded, of which 60% were due to 
drowning, mainly while the animals were traversing thin ice in 
autumn, 38% due to predators, and 2% due to other reasons. Adult 

^^'in the Pri-Amur region an excessively dry summer or an excessively rainy one is 
unfavorable for elk since availability of food is lower (Abramov, 1949). 



407 

animals drown most often since having broken through the ice they 
find it difficuh to climb out of the hole. The maximum number of 
elk drown in years of prolonged obstruction by ice floes in the 
period of autumn-winter migrations (V.P. Teplov). In the southern 
Ural deaths have been recorded for elk attempting to swim through 
floating ice in the Beluya River (Kirikov, 1952) and animals caught 
in the high waters of the Ural River (Pashkov, 1958). When the 
Rybinsk reservoir was flooded in the spring of 1941, over 100 elk 
died because debris obstructed their passage (Turov and Kuznetsov, 
1948). In Buzuluk Bor one elk drowned by falling into a pit in a peat 
bog (Teplov, 1948). Once a female elk swimming with small calves 
drowned (Peterson, 1955). 

The diseases of elk have not been studied well. The maximum 
number of references are to Siberian ulcers in the Baltic region (Yur- 
genson, 1935), Mordov Autonomous Soviet Socialist Republic and 
western Siberia (Sludskii, 1954), and Yakutia (Belyk, 1948). In some 
regions Siberian ulcers raged through the elk population (Usually 
concurrent with an epizootic of Siberian ulcers among cattle) and a 
large number died in the eighteenth and nineteenth centuries in the 
Baltic region and in 1913 and 1914 in Yakutia. In the summer of 
1950 in several regions of western Siberia carcasses of elk dead from 
Siberian ulcers were found. 
287 Instances are also known of elk suffering from cattle plague, 
necrobacillosis, emphysematous carbuncle, foot-and-mouth disease, 
and scabies. Foot-and-mouth disease has been detected among elk in 
the Leningrad district. Emphysematous carbuncle has evidently 
occurred in Buzuluk Bor (Knorre, 1953; and others). In the Komi 
Autonomous Soviet Socialist Republic domesticated reindeer have 
suffered repeatedly from necrobacillosis but only three instances of 
elk attack have been recorded (G.G. Shubin). In Karelia blood para- 
sites of family Piroplasmidae, causing hemosporidiasis, were isolated 
from the carcass of an elk (Chuzh and Bazhenov, 1951). The cause of 
elk mortality often remained undiagnosed in earlier years, for exam- 
ple in the Baltic region and in 1907 in the former Olonets province 
(Blagoveshchenskii, 1912). 

On the elk farm of Pechora-Ilych preserve gastrointestinal dis- 
eases of nursing calves are common as a result of disorders in their 
natural feeding regime. Instances are known of the death of new- 
born calves due to colibacillosis. Calves suffer from inflammation of 
the lungs, almost invariably with fatal consequences (Knorre, 1953). 
American moose suffer from brucellosis (Corner and Connell, 1958), 
tuberculosis, various diseases of the respiratory tract and other 
organs, chronic leukoencephalitis, actinomycosis, arthritis, and 



408 

malignant tumors. In captivity deaths from necrotic stomatitis have 
been estabhshed (caused by Actinomyces necrophorus) (Peterson, 
1955; Ritcey and Edwards, 1958). A mass outbreak of rabies in 
Alberta (Canada) also affected elk (Ballantyne, 1958). For over 25 
years a mysterious "moose sickness" has killed many moose in the 
USA and Canada. Its main symptoms are: loss of fear of man, lower 
mobility, uncertain movement, weakness, increased flabbiness, total 
or partial blindness, lack of response to approach, and loss of appe- 
tite. Eventually paralysis of the legs sets in and thereafter the animal 
dies. Many ticks have been noticed on affected animals. Suspicions 
about the viral and bacterial nature of the disease have not been 
confirmed (Peterson, 1955; Benson, 1957). 

Among elk inhabiting the Palearctic, 38 species of parasitic 
worms (trematodes, cestodes, and nematodes) belonging to 16 fami- 
lies have been detected. Elk is an obligate host for several of these 
species {Alcefilaria abramovi, Nematodirella alcidis, Spiculopte- 
ragia aids, and Verestrongylus alces). Some species are known parasites 
of elk throughout its range in Eurasia (Paramphistomum cervi, 
Taenia hydatigena larvae, and others), while A. abramovi and 
Parabronema skrjabini are found only in eastern Siberia and Fasci- 
ola hepatica and Parafasciolopsis fasciolaemorpha only in Europe. 
Elk in zoos serve as hosts for various helminths (Rykovskii, 1957). 

Among the most common elk helminths are Trichocephalus (in 
zoos) and Parafasciolopsis (under natural conditions). In elk 
infected with Trichocephalus the intensity of infection, according to 
available data, often reaches 5,000 or more parasites, leading to an 
extreme deterioration in the condition of the animals or even death 
(Rykovskii, 1957; Malyshev, 1958). Under conditions of intense 
infection by P. fasciolaemorpha (up to 19,000 parasites) normal liver 
functions are impaired and death may result. The intensity of infec- 
tion of elk by Paramphistomum cervi is sometimes very severe, up to 
9,000 parasites. The larvae of the latter two species of helminths 
grow in an aquatic medium. In arid years, when most small water 
holes dry up and elk are encountered around a few deep flood-plain 
water sources abounding in local intermediate hosts, the intensity of 
their infection by these helminths increases notably (Rykovskii, 
1957). 

Among helminths found in elk one should also include Elapho- 
strongylus panticola, Dictyocaulus, Echinococcus, Nematodirus, 
Ostertagia, and Moniezia. Many species of elk helminths are also 
common to agricultural animals. Several helminths occurring 

^'^Parasites in venous blood. 



409 

extensively among American moose, for example W ehrdikmansia 
cervipedes, are unknown among Russian elk (Peterson, 1955; Ritcey 
and Edwards, 1958). Among the various reasons for the mortality of 
elk, the importance of helminth-borne diseases is not generally 
great. Under natural conditions a severe infection is seen in rela- 
tively rare cases; unless the animal is very thoroughly examined, 
helminths are generally not detected (Knorre, 1953; Abramov, 1954). 

Several species of protozoans inhabit the rumen of elk (Dogel', 
1934; and others). Instances are known of wild elk infestation with 
scab mite Psoroptes equi (Uhlrich; cited from Dubinin, 1955). In 
zoos the mange mite Acarus siro has been found on elk, transferred 
to them from other ungulates (Dubinin, 1955). Among Diptera the 
naso-pharyngeal gadfly Cephenomyia uhlrichi (represented in the 
Far East by the particular form С и. kaplanovi) is prominent and 
evidently parasitizes elk throughout the range in Eurasia. This 
gadfly occurs en masse in July- August and sometimes part of Sep- 
tember. The larvae parasitize the nasopharyngeal region. They were 
present in all elk caught in Sikhote-Alin in spring. Infestation 
occurs in June to early July (Pechora basin and Sikhote-Alin). When 
the larvae exceed 100, respiration and food intake become difficult, 
molt is delayed, and severe emaciation sets in, which in rare cases 
may cause death (mainly among yearlings). These larvae grow well 
in dry soil, as a result of which their most intense infestation of elk 
occurs in the year following a dry summer (Kaplanov, 1948; Grunin, 
1957; Knorre, 1957). The larvae of the gadfly Pharyngomia picta 
infect many species of ungulates and also parasitize the nasopharyn- 
geal passage in elk. The gadflies Hypoderma alces and H. acteon are 
known parasites of elk, mainly in the western (Baltic) and eastern 
(Trans-Baikal and Sikhote-Alin) parts of the range. 

Horse flies (many species) greatly annoy elk and also act as vec- 
tors of Siberian ulcers. Their bites leave bleeding wounds on the 
hind legs of elk. Horse flies swarm en masse from mid-June through 
August inclusive (Kaplanov, 1935; Turov, 1953; and others). Other 
blood-sucking insects (mosquitoes, gnats, and biting midges) are of 
lesser importance than horse flies and gadflies, but the summer 
habitat-related dispersal of elk and their behavior greatly depend on 
the abundance and activity of pests since they affect feeding adverse- 
ly (see "Daily Activity and Behavior"). 

On elk killed in the mountains of Sikhote-Alin innumerable 
blood-sucking flies of the family Hippoboscidae and ticks (mainly 
Ixodes persulcatus) were found. Due to their attacks the bodies of the 
animals were severely ulcerated (Kaplanov, 1948). The deer louse 
Lipoptena cervi parasitizes elk in several regions. 



410 

Many herbivorous animals are competitors of elk to various 
degrees. When prolific, muskrat can pose a serious problem (they eat 
aquatic vegetation) as also deer of the genus Cervus. In Belovezh 
Forest multiplication of red deer and a high population of other 
ungulates were considered responsible for the population reduction 
of elk there at the beginning of the twentieth century. In the 1880's 
and 1890's elk in the Forest had multiplied greatly. Then, with no 
significant intervention from man, their population rapidly dwin- 
dled from 775 in 1898 to 700 in 1901, 450 in 1902, 340 in 1904, 222 in 
1907, and 58 in 1914. The living conditions for elk and other ungu- 
lates in that period in the Forest were less favorable since good tree 
and shrub foods were inadequate. Elk with signs of degeneration 
were seen — antlers without palmate portions, etc. (Kartsov, 1903; 
Vrublevski, 1912 and 1927; and others). White hare [arctic hare, 
Lepus timidus] is sometimes considered a serious competitor of elk 
(Teplov and Teplov, 1947). This is hardly correct since the hare 
strips willow at a much lower level than elk and feeds more on smaller 
shoots. 

When the elk population is extremely high, irrespective of the 
289. presence or absence of other species of ungulates, these animals per 
se can denude the winter food base. Under such conditions their 
population also begins to decrease as some elk evidently migrate, 
litter size drops, and sometimes mortality rises. In Oka preserve 
(24,000 hectares) the population density of elk, 20.2 per 1,000 hec- 
tares, had dropped by 1945 to 13.0 to 15.8 (Kozlov, 1954) and by 1951 
even to 8.8. Only after the latter level was reached did some increase 
occur (11.5 in 1952 and 12.3 in 1953); in some western sections of the 
preserve it rose to a record level of 34.4 (L.P. Borodin). In Priok- 
Terras preserve (4,800 hectares) the average density from 1952 to 1955 
was 20 to 25 elk per 1,000 hectares, which dropped to 16 or 17 in the 
next two years (Aleksandrova, 1957). In both of these preserves there 
were almost no serious ungulate competitors since sika deer (Oka 
preserve) and European bison (Priok-Terras preserve) were too few 
in number to be considered. 

Similar phenomena have been observed in other parts of the elk 
range. Furthermore, depending on local conditions, i.e., dispersal of 
winter and summer habitats, state of food (especially of winter food), 
height of snow cover, presence of competing species and predators, 
protective cover, places of escape from enemies and blood-sucking 
flies, etc., the critical population level of elk may vary from 1 to 5 to 
15 to 20 animals per 1,000 hectares. 

The ability of preserves to retain elk revolves around some defi- 
nite stages of forest "succession". For example, in the first few years 



411 

after the destruction of forest (by fire, as a result of felling, or prolif- 
eration of pests), such an area can only serve as summer pasture for 
elk. Later, depending on the availability of shoots of aspen, birch, 
goat willow, and pine undergrowth, burned over and felled sections 
become excellent winter pastures of elk, especially if an old pine 
forest section lies nearby in which the animals can find shelters. 
This situation will obtain for 10 to 20 years (the lesser duration in 
the south and the longer in the north where forest grows more 
slowly), after which the forest begins to age for elk and not only the 
amount of accessible winter food reduces, but also its quality 
deteriorates. 

In Arkhangel'sk district felled sections in the first ten years served 
only as summer pastures for elk and were not used in winter; they 
became excellent winter habitats for this species roughly from the 
twentieth year when protective conditions in them improved and 
tree food became abundant (Leble, 1953). After 40 years the ability of 
these areas to hold elk fell as food reduced. In Ontario (Canada) 
burned-over sections in relatively early stages of reforestation, i.e., 
up to 15 years, are considered most favorable for elk from the view- 
point of food availability. Their capacity increases up to 16 elk per 
1,000 hectares. At the age of 16 to 30 years the capacity of a rejuvenat- 
ing forest is assessed at 8 elk per 1 ,000 hectares, dropping to 4 at the 
age of 31 to 45 years, and to 1 after 45 years (Cringan, 1958). In 
highly snowy taiga regions the population dynamics of elk is deter- 
mined not so much by the overall food resources of a given locality, 
as by the reserves of tree and shrub foods along rivers which 
constitute winter foods for elk (Knorre, 1953). 

It is highly probable that intense changes in the northern bound- 
aries of distribution of elk and in their population in several 
regions, observed time and again in the Soviet Union, were caused 
not only by the direct extinction and subsequent colonization of elk 
from adjoining regions where they were preserved, but also by 
changes in food availability of host areas due to forest fires (Pechora 
basin and western Siberia) and mass felling of forests (central 
regions of the European part of the USSR and central Volga basin). 
Climatic changes in this case were only secondary in importance,^^^ 
290 although they might have exerted some influence on this process 



"'The dispersal of elk northward is sometimes explained by the warming up of 
the climate in the Holarctic region (Peterson, 1955; and others), but if we remember 
that there should be an increased quantum of precipitation, including snow, as a 
result of the more intense circulation of warm moist air masses, the untenability of 
this assumption becomes evident. 



412 

(for example, promoting forest fires in drought years and preventing 
them in wet years, simultaneously hindering or promoting forest 
restoration). Thus Wrangel (1841, cited from 1948 ed., p. 280) in 
describing his 1822 summer journey into the lower course of the 
Kolyma points out for one of the left hand tributaries — the Filip- 
povka River (entering the Kolyma below Bol'shoi and Malyi Anyui): 
"The Filippovka valley was formerly well known for many sokhati 
or Siberian elk. The great forest fire of 1770 drove them away, but 
later, as the forest rejuvenated they reappeared and multiplied so 
much within a short period that by 1812 rare was the hunter who did 
not bag half a dozen sokhati. Ten inhabitants of Panteleevka village 
caught 70 of them. One cannot state for certain the reasons, but after 
this event, sokhati again disappeared, and it is my understanding 
that this hunt was among the last of the lucky strikes. During our 
residence the appearance of sokhati in the Kolyma region was 
already considered a rarity and described as a bygone epoch in hunt- 
ing. On the Omolon and south of it elk were still resident but in 
small numbers." Buturlin (1913), who visited the Kolyma River basin 
in 1905, remarks that elk were old inhabitants in its delta; they were 
absent there in the 1870's and 1880's but reappeared subsequently. 
The composition of elk populations has been better studied than 
for most other wild ungulates. Among newborns the sex ratio varies 
highly, but on the average there is evidently some preponderance of 
males although the picture may be quite the opposite in some individ- 
ual regions and years. In 50 elk litters from different regions of the 
country males represented 56% and females 44%, while in Buzuluk 
Bor (17 calves) males constituted only 30% but in Pechora-Ilych 
preserve (14 calves) 79% (Knorre, 1949). Later, a large amount of 
material from Pechora-Ilych preserve revealed that males constituted 
62% and females 38% (V.P. Teplov). At places where elk are not 
killed the sex ratio in the adult population usually evens out to 1:1, 
but quite often favors females. This is evidently due to the higher 
mortality rate of males as a result of severe emaciation during rut, 
greater mobility and evidently less caution, and death resulting from 
fights over females. ^^*^ In Lapland preserve among elk attacked by 
bear males constituted 67% and females 33% (Nasimovich and 
Semenov-Tyan-Shanskii, 1951). In Pechora-Ilych preserve among 19 
elk which died for various reasons, males were somewhat more 
numerous (10) and in Buzuluk Bor numbered 8 among 12 dead 
(Teplov, 1948). 

^*°Of 12 elk dead in Buzuluk Bor for various reasons, 5 were the result of injuries 
received by males in combats over females (Teplov, 1948). 



413 

In Lapland preserve among elk older than one year the sex ratio 
in the period June-December is close to 1:1 (97 females and 96 
males; Semenov-Tyan-Shanskii, 1948). In Pechora-Ilych preserve 
the sex ratio of 2,330 elk older than one year was established in yards 
from 1938 to 1948; males comprised 52% and females 48% (Zharkov, 
1952).^^' In Priok-Terras preserve winter counts from 1953 to 1955 
showed that males among elk older than one year varied from 38 to 
48% in different years and females were correspondingly 62 to 52% 
(Aleksandrova, 1957). In the Lithuanian Soviet Socialist Republic 
similar counts have shown that males averaged 42% and females 58% 
(V.A. Bergas). In ten regions of the- European part of the USSR in the 
winter of 1943 to 1944 males averaged 42% (39 to 45%) and in the 
winter of 1945 to 1946, 43.8% (Danilov, 1949). 

Selective hunting may strongly perturb the natural sex ratio. For 
example, at the beginning of the first decade in the hunting farm 
near present Krasnoe Selo (Leningrad district), there were only 60 
291 males to 294 females. In other words, males represented only 17% of 
the total number of adult elk. As a result mating of females with 
juvenile males increased; such unions were evidently not always 
productive as off-spring were few (6.3%) and instances of degenera- 
tion observed time and again (Sablinskii, 1914). 

The number of calves in a population is higher at the beginning 
of summer and lower in winter. Elk mortality is highest in the first 
one or two months of life (Knorre, 1953). By mid-winter calf mortal- 
ity in the European part of the USSR has reached 50% (Danilov, 
1949). In Pechora-Ilych preserve about 35% calves die by September- 
October and up to 60% in the first ten months (Teplov and Teplov, 
1947). In Pechora-Ilych preserve calves in the population from June 
to August of different years have varied from 18 to 25%, on the 
average 21% (Teplov, 1948), and in yarding sites in winter from 7 to 
23% (Zharkov, 1952). Young elk are few after winters of severe snows. 

Elk counts in ten regions of the European part of the USSR 
revealed that the number of calves in the winter population of 1943 
to 1944 varied from 25 to 33% (average 27%) and was 23.9% in the 
winter of 1945 to 1946 (Danilov, 1949). In the Tatar Autonomous 
Soviet Socialist Republic juveniles in winter in different years (1944 
to 1950) constituted 26 to 29% (Aspisov, 1955) and in the Lithuanian 
Soviet Socialist Republic 17 to 22% (V.A. Bergas). In Lapland pre- 



^*'During winter calculations, which provide the bulk of data for elk, the possibi- 
lity should always be borne in mind that less experienced observers could mistake a 
number of males as females since at that time of year most do not bear antlers. 



414 




Fig. 89. Tracks of female elk with two calves (animals moving away from observer). 
Lapland preserve. March, 1951. Photograph by O.I. Semenov-Tyan-Shanskii. 

serve the average for June to December was 21% (Semenov-Tyan- 
Shanskii, 1948). 

Knorre (1953) considers 20% or more offspring in an elk popula- 
tion high, 15% average, and 10% or less low. In Canada this annual 
increase in elk population has on occasion exceeded 20% with an 
average of 16% (Peterson, 1955). 

Field characteristics. Elk are not difficult to distinguish among 
other ungulates. Even young elk are readily distinguished from 
young of other deer by their long legs and absence of spots. 

In winter, when males have no antlers (November to April), at a 
distance sex identification of elk from external appearance is diffi- 
cult, since such characters as the usually smaller body size, lesser 
development of the dewlap and "beard," and slightly different pat- 
tern of coloration around the tail, etc. in females are not readily 
discerned. One must also take into consideration the group 



415 




г^'Г i^p/ffrapTT^r.^rx; 






292 Fig. 90. Track of elk in sand (without imprints of lateral digits). Lapland preserve. 
June, 1950. Photograph by O.I. Semenov-Tyan-Shanskii. 

composition — adults versus calves. The urine stream in females falls 
between the imprints of the rear legs and in males in front of them. 
In females the hoof is prolonged with very narrow and sharp ends; 
in males it is usually larger, broader, and circular with blunt ends. The 
292 tracks of males and females are therefore somewhat different but 
such differences are difficult to discern in soft snow. 

The track of a moderate-sized male elk on the ground, excluding 
the imprints of lateral digits, is 152.5 mm long and 127.5 mm wide; 
in females the corresponding values are 140 and 101 mm (Buturlin, 
1934). In large elk the track length can be as much as 175 mm. In 
yearling elk it is roughly 100 to 120 mm. The track of an average- 
sized male elk is invariably larger than that of a cow. Elk usually 
follow a beaten trail. 

The direction of movement of elk on soft snow is best judged 
from the scatter of snow in front of the track and the greater com- 
pactness of the front wall of the "track impression"*, since the 
animal drags snow while shifting its legs (dragging in and dragging 
out of each track) and from other clues of movement (around bushes, 
toward trees, etc.). 

Adult elk stand 160 to 216 cm, more often near 175 cm, at the 



^Literally "glass [cup] of the track". 



416 



Fig. 91. Winter feces of elk (natural size). Moscow district. Sketch by T.L.Savranskaya. 

withers. Calves in the first year are no more than 135 cm and in the 
second year 155 cm. The pace of an adult elk is 60 to 90 cm, which 
lengthens to 160 to 200 cm while trotting, and up to 3.5 m in 
jumping. 

Elk lie on the snow without pawing it. The bed is kidney-shaped 
and 100 cm X 170 cm, 90 cm x 140 cm and 60 to 65 cm x 100 to 1 10 cm 
for males, females, and calves respectively. The depth of the bed into 
the snow surface may be 30 to 40 cm (Semenov-Tyan-Shanskii, 
1948). Since elk sometimes stand only to lie down again a short 
distance later, the number of beds is often more than the number of 
elk in a group. 

Winter and summer feces are in the form of oval pellets (4.0 to 5.0 
cm X 1.5 to 2.0 cm; those of calves are not longer than 2.5 cm). In 
winter they are much firmer, with remnants of chewed trees visible 
in them; in summer they have an oily sheen, are black, soft, cohesive, 
and sometimes bear traces of semidigested grass. Widely recurring 
293 statements in literature that female elk feces are invariably in the 
form of long pellets and those of males rounded are totally false 
(Knorre, 1953). In transitional seasons, especially spring, elk feces 
are often of a semifluid consistency, resembling those of bovines. 

In summer pastures elk often nibble tops of tall plants (willow 
herb [=fireweed] and others); at places of feeding in the water, bits 
of Nuphar and water lily rootstocks, their leaves, etc. are often seen 
floating. Elk scrape tree bark with their incisors, leaving behind a 
chisel-shaped groove, or tear long strips by gripping the edge with 
the lower incisors and moving away from the tree. Sometimes bark is 
stripped to a height of 100 to 350 cm from the ground. Adult elk 
tear off the shoots of trees (first breaking them) most often at a height 
of 250 to 270 cm above the ground. 

In rubbing the velvet off their antlers, and later right up to the 
moment antlers are shed, elk often use them to break young (up to 8 
to 10 cm diameter) trees and twist and reduce them to splinters. In 



417 




Fig. 92. Summer feces of elk (natural size). Moscow district. Sketch by T.L. Savranskaya. 



addition they often maul and uproot bushes. Males in rut dig pits 
with the hooves of their forelegs to depths of 25 to 30 cm and up to 
80 to 120 cm wide. The urine of the male in these places is highly 
pungent. 

The voice of elk is most often heard in the period of rut when 
males moan (see "Reproduction" above) and females (often during 
mating) emit a sound similar to a snore or neigh. Calves emit a very 
soft moo. 

Economic Importance 



Among elk products, the meat is of utmost value. Its quality is super- 
ior at the end of summer to early autumn when a single elk can yield 
30 kg of internal and subcutaneous fat (Kaplanov, 1948). After rut 
male elk lose up to 17% of their former weight (in the period of rut 
their meat is less edible) and later recover only slightly before the 
onset of winter. Females remain well-fed up to December through 
early January; barren females retain their weight slightly longer. Elk 
weight is lowest at the end of winter and in spring. 

The body weight of an adult elk is most often 300 to 400 kg (for 
details, see section "Growth, Development, and Molt," and Table 3 
above) but at places where elk are hunted intensely their weight 
averages a little over 200 kg. In autumn the meat yield varies from 
60.0 to 62.5% (Buturlin, 1934; Knorre, 1939; Popov, 1939) to 64 to 65% 
of body weight; in barren females it possibly goes up to 70%" ~ (Lik- 

^'**Tfie meat yield of 69.5 to 75.0% quoted by Kaplan (1948) for Ussuri elk is an 
exaggeration. 



418 

hachev, 1955). At the end of winter the meat yield is much less since 
some elk during the period of estrus and in the following winter lose 
up to 80 kg (Knorre, 1953). The maximum meat yield of Pechora elk 
is 300 kg (E.P. Knorre). 

Taking the weight of a European elk without antlers as 100%, the 
head (without antlers) constitutes 6.25% of total weight, fresh hide 
with lower parts of legs 10.75%, stomach and intestines (with con- 
tents) 17.0%, and carcass with liver, heart, and kidneys 62.5% 
(including blood, which accounts for 3.5%) (Buturlin, 1934). In 
294 Yakutia it was established that the head weight without antlers con- 
stitutes 7.15% of total weight, fresh hide 11.75%, stomach and intes- 
tines 18.25%, and carcass 60.1% (Popov, 1939). Elk fat is of a greater 
fluid consistency than that of cows and has a good flavor. When 
animals are raised in captivity, their milk can also be used (see 
"Growth, Development, and Molt"). It is high in fat content and has 
a taste similar to the cream of cow's milk but without a salty- 
astringent aftertaste. The maximum milk yield in a season is 430 
liters per elk but fat content corresponds to 1,290 liters of milk from 
local cows. One liter of elk milk can yield 1 10 g butter (E.P. Knorre). 

In some regions local people also consume the blood of elk. The 
cecum is stuffed with congealed blood admixed with bits of meat 
and fat and sausages made from it (Amur and Sikhote-Alin; 
Abramov, 1954). 

The thick strong hide is converted into high-quality chamois. 
Local people also used it in the recent past for making fur coats 
(with the fur outside; heavier than that of goat and deer), dresses, 
parkas (with fur inside), caps, winter footwear, mittens, etc. Elk 
hides are often used as bedding even today. Belts, straps, harnesses, 
and knitting bags are other products made from hides. The skin of 
the legs is stitched or glued to skis and used for soles of high boots. 
In summer flaws reduce the value of hides. The hair is utilized in 
mattress stuffing and furniture padding. Elk chamois is better than 
deer chamois in quality and in the eighteenth and nineteenth cen- 
turies was converted into breeches and other garments for battalions. 

Antlers are used as trophies, in making knife handles, etc. In the 
past a medicinal meat-jelly was made from the horns and considered 
curadve for tuberculosis (Turkin and Satunin. 1902 and Solov'ev, 
1922). A good quality glue can be obtained from hooves and tendons 
are used as thread (Buturlin, 1934). 

In the early 1920's about 70,000 elk were caught annually 
(Solov'ev, 1922). A much higher figure has been quoted for the end 
of the nineteenth century (Turkin and Satunin, 1902), but such 
figures are purely speculative. In the eighteenth and early nine- 



419 

teenth centuries elk as an object of hunting was evidently much 
more irriportant and the well-being of local peoples at many places 
depended on the success of the hunt. In the seventeenth century, and 
probably even later, in the Sosva basin an elk cult developed among 
the local populace (Bakhrushin, 1935). According to data recorded 
in 1926 and 1927 for the pri-polar north of the USSR (data for 
Yakutia destroyed), 5,564 elk were caught in 1925 in regions of the 
Far North. The total catch of elk in the country in the early 1930's 
was placed by Buturlin (1934) at 10,000 which, according to him, 
exceeded the herd growth at many places. 

In the hunting season of 1960 to 1961 in the Soviet Federal 
Socialist Republic licenses were issued for killing 18,000 elk (N.V. 
Eliseev). Actually, throughout the USSR no less than 35,000 elk are 
caught nowadays, but even with this number elk reserves are not 
thoroughly exploited (see "Population"). In many regions licensed 
hunting of elk should be increased, but poachers simultaneously 
discouraged. The permissible norm of hunting should be 15% of the 
total population (Knorre, 1953). Permission should be granted to 
hunt not only males but also females, but males should predominate 
roughly 2:1. When males are hunted in a higher ratio, the number of 
barren females increases notably. 

Hunting elk at the end of summer (August 15 to September 15) is 
futile since the meat goes bad quickly at that time of year. Hunting 
elk in the period of rut is likewise foolish since the meat of males at 
that time is less edible; furthermore, hunting frightens the animals 
and barrenness subsequently increases (Peterson, 1955; and others). 
The best hunting period is from November 1 to December 1 
(depending on the period of termination of rut and allowing two to 
three weeks thereafter for males to recoup somewhat, for a total 
duration of not more than LO to 1.5 months). 

At present the greatest numbers of elk are caught with the help of 
295 dogs. Hunting with dogs is generally done in autumn and early 
winter on rough trails when the snow is still sparse. The hunter 
releases a laika dog (sometimes two or three) on the fresh tracks of an 
elk, the dog stalks the animal, and the hunter's approach to the 
quarry is thereby simplified. This type of hunting is practiced exten- 
sively throughout Siberia and the Far East. Sometimes the dog is 
kept leashed and the hunter moves along with it (Karelia, Irkutsk 
district, Pri-Amur, and others). During this maneuver the dog 
should lead the hunter to the elk without raising its voice; the hun- 
ter's job is thus doubly simplified. 

In autumn and early spring, when the snow is not deep (up to 20 
to 30 cm), especially during a fresh snowfall, elk are hunted on foot 



420 

rather than on skis. In the past hunters rode on sledges (when the 
animals were less frightened) or on horseback (the Sayans) but these 
methods have now been mostly discarded. On skis hunters are rela- 
tively poorly hidden from elk since skis make considerable noise. 
Most elk in the European part of the USSR are hunted from places 
of concealment and by chasing (for which purpose a few hunters 
band together). This type of hunting is practiced in Siberia also. 

Elk are also caught by organizing drives; sometimes flags are 
used but not all elk are frightened by them. In Siberia and Komi 
Autonomous Soviet Socialist Republic during autumn elk are often 
killed by shooting them in the course of their seasonal migrations. 

Most other methods of catching elk are either banned or have 
been discarded. Among such methods are catching elk by driving the 
animals into an enclosure made on deep snow and surface crust. In 
the pre-Revolutionary period and the first few years after the Great 
October Revolution this method prevailed. Nowadays it is employed 
exclusively by poachers, mainly in the least supervised parts of the 
elk range. Chasing an elk is possible only when the snow is very 
high (a meter or more) but even then two or three men on skis often 
have to chase an animal for several days in relays before coming 
within firing range. If there is an ice crust on deep snow, hunting is 
greatly facilitated and often ends quickly; on crusted snow laika 
dogs are sometimes used to help arrest elk movement. At the end of 
winter elk become greatly emaciated and their meat dry and devoid 
of fat. Sometimes animals which elude the chase become overheated 
and later die. 

There is no hunting of elk in the summer season. Formerly, elk 
at that time of year were often caught on water (sought by elk to 
escape blood-sucking flies and to feed on aquatic vegetation). Either 
an ambush was set up or a stealthy approach made from the bank or 
a boat. Elk are also hunted at salt licks (Sikhote-Alin) with the aid of 
dogs, but in summer even an experienced dog finds it difficult to 
restrain an elk. Autumn hunting from rafts, practiced mainly by 
privileged hunters in pre-Revolution Russia, has been discontinued. 
Hunting with a birch tube ("decoying") in the period of estrus, the 
hunter imitating the voice of a cow to lure a male, is also on the wane. 
This type of hunting ("hunting the moan") is largely a form of pure 
sport, since the meat of males at that time is hardly suitable for human 
consumption. 

In pre-Revolutionary Russia various methods of trapping elk 
were widely employed, mainly by setting up long (up to 10 to 15 km) 
fences with exits in them where pits were dug, crossbows held at the 
ready, "runners" set (a log with a knife embedded in it to rip the 



421 

animal's underbelly), and large traps laid. Fences were constructed 
at places elk passed through during seasonal migrations. A large 
number of elk falling into pits died for nothing since the pits were 
not supervised properly (the Urals, western Siberia, Yenisey basin, 
Trans-Baikal and Sikhote-Alin). Crossbows were also used at salt 
licks, on tracks, etc. This type of elk hunting was brutal and gener- 
ally dangerous, and hence has been banned. Nevertheless poachers 
practice it even now, not only in Siberia but also in the European 
part of the USSR. Usually metallic snares and sometimes small 
guide fences are set up (Komi Autonomous Soviet Socialist Repub- 
lic; O.I. Semenov-Tyan-Shanskii). 
296 Experience gained in Pechora-Ilych preserve has shown that fen- 
ces can be successfully used for catching elk in small enclosures. Part 
of the elk corraled are killed and part set free. The maximum 
number are trapped in such enclosures during migrations (Shubin 
and Yazan, 1958). 

When the elk population is high, saplings and bushes in the 
forest and forest nurseries incur great damage. ^^^ In the European 
part of the USSR and also Scandinavia young pines (more often not 
older than 20 years) are damaged most. In the southern parts of the 
range many valuable species such as oak are also damaged by an 
excessive elk population, but only at places where pine are absent or 
few. Damage by elk to aspen, willow, mountain ash, alder, birch, 
juniper, etc. is of less significance to the forest economy since these 
species are less valuable. In the European part of the USSR the zone 
in which pine are severely damaged by elk extends from the Baltic 
region and the upper course of the Dnieper to the Ural River. Dam- 
age is maximum at places where a single elk has for itself an average 
of less than seven hectares of young pine forests (Dinesman, 1957). 
The critical form of damage is breaking of tree tops. Observations in 
Sweden have revealed that when the elk population density is 
extremely high (up to 30 animals per 1,000 hectares), up to 80% of 
the pine forest is damaged and 10 to 50% of the trees perish. Damage 
is maximum at places where deciduous species are intermixed with 
pine forests and higher in less dense forests (Westman, 1958). 

Various methods have been implemented to protect valuable 
forests from elk: planting less valuable species along forest roads, 
feeding elk hay and cut aspen (in most regions elk consume hay 
poorly), fencing individual forest sections, applying oily substances 
to trunks to repel the animal, wrapping trunks with nylon, etc. 
(Westman, 1957; Dinesman, 1957; and others). None of these meas- 

Also see section "Population". 



422 




Fig. 93. Aspen with bark peeled by elk. Moscow district, 1949. Photograph by A.G. 

Bannikov. 



297 



ures attain the objective. They are either too costly or insufficiently 
effective. The only rational measure is the regulation of elk and a 
more intense utilization of their stocks. 

Elk rarely invade cultivated fields; at places, however, elk in 
spring have damaged winter crops, oats, and melon fields (Voronezh 
district). 

In the past various deformities in cattle calves were often ascribed 
to the mating of cows with elk. This is completely impossible 
since these animals belong to different families. 

Their large size, great strength, ability to easily negotiate areas 
cluttered with windfallen trees, hummocky or swampy areas, and 
high snow cover prompted people long ago to attempt domestica- 

^^''This possibility was endorsed quite recently by Manteifel' (1934). 



423 

tion of elk. Domestication was facilitated by the relatively easy feed- 
ing of newborns, their tameness, and the abundance of winter food 
for elk. Later, their early maturation also became known; this, like 
their fertility, is better than that of domesticated cattle. Efforts to 
tame elk and harness them in the Soviet Union are known from the 
eighteenth century. ^^^ Right up to the end of the 1920's, however, 
such attempts continued to be random although essayed several 
times. Attempts to tame elk were better organized in Berezinsk 
reserve area (Belorussia), Yakutia experimental station, Serpukhov 
farm, and Buzuluk Bor. But this work has been carried out com- 
prehensively and along proper lines only in Pechora-Ilych preserve 
where, in the early 1940's, an experimental elk farm was organized 
under the guidance of E.P. Knorre, formerly engaged in the domesti- 
cation of elk in Buzuluk Bor. Elk reproduced well in the experimen- 
tal farm. Several dozens have been raised and many tamed and used 
as pack, transport, and draft animals. A test run in harnessing elk 
was successfully conducted in winter over a distance of 240 km; 
during the test run they were fed branches and potatoes (G.G. 
Shubin). 

Elk work best at a slow pace. Their most promising use is as pack 
animals and partly as transport animals in forests without roads for 
hunters, search parties, etc., especially at places where there are no 
lichens and the maintenance of reindeer is unprofitable. Elk are 
more successfully used in autumn and the first half of winter. Cas- 
298 trated males (bulls in rut are dangerous) are most convenient for this 
purpose. These animals are incapable of working in the heat of 
summer. In the latter half of winter they become highly emaciated 
and less suitable for movement in thick snow cover. Elk also cannot 
be gainfully employed in spring when molt occurs. Thus the total 
duration of the effective utility of elk is evidently limited to less than 
40 to 50% of the year. 

The packload of yearling elk is up to 50 kg and on sledges up to 
100 kg; corresponding values in the second year are 75 and 200 kg, 
and in the third year 100 and 300 kg, respectively. The daily run of 
yearlings should be restricted to around 10 km, in the second year to 
20 km, and in the third year to 30 km (Knorre, 1953). The maximum 

^^^Skalon and Khoroshikh (1958) think that the domestication of elk and their use 
as transport animals was in vogue even from the Neolithicperiod and was particularly 
well developed in the Iron Age, declining only later. This supposition is not sup- 
ported by adequate data. 

^^^Buturlin (1934) lists several such studies. 

^"Experiments were also conducted in Kostroma reserve zone, Tyumen, Novosi- 
birsk zoo, and in Sikhote-Alin preserve. 



424 




297 Fig. 94. Elk as pack animals. Pechora-Ilych preserve. October, 1952. Photograph by 

O.I. Semenov-Tyan-Shanskii. 

packloads are 135 kg and on sledges 500 kg (Popov, 1939) or even 
considerably more (E.P. Knorre). If trained from an early age, elk 
can be made to move quickly (E.P. Knorre). 

Recently elk raised in Pechora-Ilych preserve were set free in an 
experiment throughout the summer. By autumn of 1958, 19 of the 22 
elk released returned to the pen (the rest possibly died or were caught 
by poachers). Elk of the second and third generation raised in captiv- 




Fig. 95. Elk harnessed to a sledge. Pechora-Ilych preserve. December, 1953. Photo- 
graph by O.I. Semenov-Tyan-Shanskii. 



425 

ity were far easier to train than the first generation (E.P. Knorre). It 
would be premature to draw decisive conclusions just now on the 
profitability of raising elk. But it is quite clear that farms designed 
for selective catching of elk during their migrations should be self- 
supporting. 

Elk remain year-round in many preserves. Some play a promi- 
nent role in restoring the elk population (Pechora-Ilych, Konda- 
Sosva, Altai, the Sayan, Sikhote-Alin, Oka, and others). 

Within the Soviet Union experiments to reacclimatize elk have 
evidently only been conducted in Belovezh Forest where a nursery 
was set up and 10 elk from it set free in 1938 and 1939. By 1946, of 
these elk only three had survived (Sablina, 1955). It would probably 
be more useful to acclimatize elk in Kamchatka (Buturlin, 1934) 
where they would not encounter other competing species of ungu- 
lates. Serious attention should be given to the reacclimatization of 
elk in the western and central parts of Altai where they were extermi- 
nated in the nineteenth century. It would obviously be less useful to 
release elk in the northern Caucasus (Buturlin, 1934). (A.N.) 

GENUS OF REINDEER* 

299 Genus Rangifer H. Smith, 1827 

\775. Rangifer. Frisch. Natur.-Syst. d. vierfiis. Thiere, p. 3. The 
name does not meet nomenclatural requirements. 

1821 . Rangifer. H. Smith. Griffith's Cuvier Animal Kingdom. 
Mamm. Syn., p. 304. According to a resolution of the Interna- 
tional Committee on Nomenclature (resolution No. 91), this 
should be used as the generic name. Cervus tarandus Linn. 

\S27. Tarandus. Billberg. Syn. Faunae Scand., vol. 1, p. 22. Taran- 
dus lapponum Billher g=Cervus tarandus Linn. 

ISib.AchUs. Reichenbach. Naturgesch. Saugeth., vol. 3, p. 12. Sub- 
stitute for Tarandus Billberg. (V.H.) 
Rangifer are telometacarpal deer of moderate size, relatively 

slender build, with a fairly long trunk and neck and relatively short 

legs. Head proportionate in size. Tail short. 

Entire metapodial region massive. Hooves of middle digits large, 

broad, and blunt. Hooves of lateral digits quite large and set low. 
In general proportions the skull corresponds to that of red deer 

(Cervus) or is only slightly more elongated. Overall length of skull 

over twice that of malar width. Facial part of skull relatively high in 

*Literally, "northern deer" — Sci. Ed. 



426 

region of commencement of nasals. Distance from front tooth to 
anterior end of premaxilla more than length of upper tooth row. 
Premaxilla of moderate length but longer than upper tooth row. 
Anterior end of premaxilla with a small pit. Nasal processes of 
premaxillae reach nasals. Nasal opening of normal dimensions, its 
length not exceeding that of the nasals. Nasals not shortened and 
longer than upper tooth row. Orbits tubular, project laterally. 
Lacrimals relatively narrow and elongated. Preorbital depression on 
lacrimal long and distinct but not deep or sharp in outline. Ethmoid 
pit large. Antler pedicels of frontals set upwards. Rear of vomer high 
and divides choanae into two separate" portions. Tympanic bulla 
relatively small and auditory tube long. Lower jaw elongated and 
symphysis roughly equal to length of lower tooth row. 

LIpper canines present and well developed in males, but very 
small in females. Upper molars relatively small, width less than 
height, and size of premolars hardly differs from that