Mammals

of the
Soviet
^
^

Union

VOLUME
Rut lb

VG.
N.

Heptner

E Naumov

.-

'^-

^-s^--

The

present book, of the series

Mammals of
is

the Soviet Union: Sirenia

and Carnivora,

the

fourth volume to be published in English.

Previously Volume
Perissodactyla,

(Hyenas and Cats)

in 1988, 1992,

Artiodactyla and Volume Part 2 Carnivora and Volume Part

I

II,

II,

Pinnipeds and Toothed Whales were published

and 1996 respectively. Sea Cows, Wolves and Bears constituted the second
in the series to appear.

The

original Russian

book

is

so large (982

pp.) that its publication in the English

translation format would have resulted in a book of 1600 pages, which was considered too unwieldy. Therefore, it was decided to publish Volume II, Parti Sirenia and

Carnivora, in two roughly equal parts.

first (Part la)

The

consisted of the Steller's sea

mammal once found only around the Commander Islands in the western Bering Sea, and the Russian species of the wolf and bear families. This second half (Part lb) comprises the various
cow, the now-extinct marine

members of the weasel

family, plus the

introduced American raccoon.

As indicated in the foreword to the English edition of the Volume I, the senior author and editor of the series, Dr. Vladimir
Georgievich Heptner, died in 1975, and did not complete the projected series. However,

work continues, and Russian volumes on baleen whales, lagomorphs and dipodid
rodents are

now

published.

A final
Volume

point: since the appearance of

II, Part 2, the Soviet Union has disappeared. However, the series was written in the context of the former "union of

republics,"

and

edit the text to

it would be confusing to conform to the current

re-

political reality.

Mammals

of the Soviet Union

Volume

II

Part lb

MAMMALS OF THE
In Three

SOVIET UNION

Volumes

Edited by

V.G. Heptner and N.P.

Naumov

Vysshaya Shkola Publishers Moscow, 1967

Mammals

of the

Soviet Union
Volume

II,

Part lb

CARNIVORA
(Weasels; Additional Species)

V.G. Heptner, N.P. Naumov, P.B. Yurgenson

A. A. Sludskii, A.F. Chirkova and A.G. Bannikov

Illustrators

A.N. Komarov and N.N. Kondakov

Scientific Editor

Robert

S.

Hoffmann

Smithsonian Institution Libraries

and

The National Science Foundation

Washington, D.C.

2001

Mlekopitayushchie Sovetskogo Soyuza In Three Volumes V.G. Heptner and N.P. Naumov, editors

Vysshaya Shkola Publishers

Moscow, 1967
Translators: Bolos

Abdul Malek Botros, Hamed Tantawi, Hosni Ibrahim Youssef, Ali Abdul Moneim Moussa General Editor: Nawal Mehallawi, Al-Ahram Centre for Scientific Translations, Cairo, Egypt

2002 Amerind Publishing Co.

Pvt. Ltd.,

New

Delhi

Library of Congress Cataloging-in-Publication Data (Revised for volume 2, pt. lb)

Geptner, V.G. (Vladimir Georgievich), 1901-1975

Mammals

of the Soviet Union.

Translation of: Mlekopitaiuschie Sovetskogo Soiuza.

Bibliography:
Vol. 2,
pt.

v. 2, pt. la, p.

lb has index.

Supt. of Docs, no.: Si 1.2

Ar 7/2

Contents: vol. 1. Artiodactyla and Perissodactyla vol. 2 pt. la. Sirenia and Carnivora (Sea Cows: Wolves and
Bears)

Carnivora (Weasels; Additional species) Carnivora (Hyaenas and cats). Soviet Union. I. Nasimovich, A. A. II. Bannikov, 1. Mammals AndreT Grigor'evich. Ill Hoffmann, Robert S. IV Title.
vol. 2. vol. 2.
pt.

lb.

pt. 2.

QL728.S65G4713

1988

599.0947

85-600144

Published for the Smithsonian Institution Libraries by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110001
Printed
in India at

Baba Barkha Nath

Printers,

New

Delhi.

'

Table of Contents
Vol.
II,

Part lb

Family Mustelidae (Martens, Weasels, etc.) Key for Identification of Genera in the Family Mustelidae
Subfamily Mustelinae Genus Martes Pinel, 1792 (Martens)

735

746 749 749

755

Key

for Identification of Species in the

Genus Martes
Martes {Martes) zibellina (Linnaeus, 1758)
(Sable)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation

Biology
'

Practical Significance

757 759 759 770 770 787 797 823

825

Martes {Martes) martes (Linnaeus, 1758) (Forest Marten)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation Biology

Practical Significance

826 826 836 836 847 850 872

Martes {Martes) foina Erxleben, 1777 (Stone Marten)

Diagnosis
Description

Systematic Position

874 875 876 882

Geographic Distribution Geographic Variation Biology

Practical Significance

883 891 895 904

Martes (Charronia) flavigula Boddaert, 1785 (Kharza, or Yellow-throated Marten)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

Genus Gulo Storr, 1780 Gulo gulo (Linnaeus, 1758) (Wolverine)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

Genus Mustela (Linnaeus, 1758)

905 905 905 910 910 912 914 919 920 925 926 926 931 932 940 943 953 954

Key

for Identification of Species in the

Genus
962
965 966 967 972 972 975 985 995

Mustela Mustela {Mustela) nivalis Linnaeus, 1766

(Weasel)
Diagnosis
Description

Systematic Position

Geographic Distribution

Geographic Variation
Biology
Practical Significance

Mustela {Mustela) erminea Linnaeus (Ermine)

Diagnosis
Description

1758

Systematic Position

Geographic Distribution

995 997 997 1003 1003

Vll

Geographic Variation
Biology
Practical Significance

1008

1016 1029 1030

1031

Mustela (Mustela) altaica Pallas, 1811

(Solongoi)

Diagnosis
Description

1031

Systematic Position

Geographic Distribution Geographic Variation Biology

Practical Significance

1036 1037 1043 1047

1051
1051

Mustela {Mustela) sibirica Pallas, 1773

(Kolonok or Siberian Weasel)

Diagnosis
Description

1052 1052 1058 1058 1065 1069 1078

1078

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

Mustela {Mustela) lutreola (Linnaeus, 1761)

(Russian, or European Mink)

Diagnosis
Description

1080 1080 1086

1088 1094 1099 1106
1107 1108 1108
1115 1117

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

Mustela {Putorius) putorius (Linnaeus, 1758)

(Forest, or Black, Polecat)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation Biology

Practical Significance

1124 1126 1132

vin

Mustela (Putorius) eversmanni Lesson, 1827

(Steppe, or White, Polecat)

Diagnosis
Description

Systematic Position

1133 1134 1135 1142

1145

Geographic Distribution Geographic Variation

Biology
Practical Significance

1158
1165 1175

Genus Vormela Blasius, 1884 Vormela peregusna (Giildenstaedt, 1770)

(Marbled Polecat, Perevyazka)
Diagnosis
Description

1176 1180
1181 1181 1188

Systematic Position

Geographic Distribution
Geographic Variation
Biology
Practical Significance

1188 1194

Genus Mellivora
(Ratel, or

Storr,

1780

1202 1206 1207

1211 1211

Mellivora capensis (Schreber, 1776)

Honey Badger)

Diagnosis
Description

1211

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

1217 1217
1222 1223 1228 1228 1228 1232 1234 1234 1242 1242 1249
1262 1281

Subfamily Melinae

Genus Meles Brisson, 1762 [=Boddaert, 1785] Meles meles (Linnaeus, 1758)
([Eurasian] Badger)

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

IX

Subfamily Lutrinae

1283 1283

Genus Lutra Brisson, 1762 [=Brunnich, 1771]

Lutra lutra* (Linnaeus, 1758) ([Eurasian] Otter)

1288

Diagnosis
Description

Geographic Distribution Geographic Variation

Biology
Practical Significance

1289 1289 1297 1305 1310 1327

Otter,

Genus Enhydra Flemming, 1822 Enhydra lutris (Linnaeus, 1758) (Sea

or Kalan)

1330 1333
1333

Diagnosis
Description

Systematic Position

1334 1343
1343 1348 1350 1365

Geographic Distribution Geographic Variation

Biology
Practical Significance

Supplement to the Order of Carnivores Family Procyonidae (Raccoons, etc.) Subfamily Procyoninae Gill, 1872 Genus Procyon Storr, 1780 Procyon {Procyon) lotor (Linnaeus, 1758) (American Raccoon)
,

1366 1366 1373 1373

1374 1374 1374 1377 1377 1383 1383 1390

Diagnosis
Description

Systematic Position

Geographic Distribution Geographic Variation

Biology
Practical Significance

Family Mustelidae Mustela (Mustela) vision Schreber, 1777 (American Mink) Diagnosis
Description

1391
1391

Systematic Position

1392 1397

*In Russian original, Lutra (Lutra)

Sci. Ed.

Geographic Distribution
Geographic Variation
Biology
Practical Significance

1399 1403 1405

1415

LITERATURE CITED
INDEX OF LATIN NAMES OF TAXA*

1416 1539

*The Russian

original also included an Index of Russian

Names, omitted

here.

491

Family of Martens
Familia

MUSTELIDAE

Swainson, 1835

These are specialized, and in part highly specialized predators of small and very small dimensions, but with several being of moderate dimensions (the smallest species in the order belong to this
family).

The hind limbs

are short, the trunk

is

strongly elongated

and

thin,

but rarely comparatively short and compact.

are digitigrade or semi-plantigrade, but several are

The limbs

plantigrade with five digits on the fore- and hind-limbs. The ventral
surface of the hind foot and forepaw are covered with hairs (as
usually, the skin of the interdigital
is,

membranes) or with naked

foot-

pads or entirely bare; the claws are of moderate length to very large.
492 In several (individual species in the otter subfamily), bare

swimming

membranes

between the digits are well-developed, the claws are greatly

reduced (Aonyx), or the digits on the forelimb are shortened and closely united with one another; while on the hind limbs they are elongated

and transformed into a hair-covered flipper {Enhydrd). The claws are not retractile and the phalanges of the digits have a normal articulating surface (the last phalanges cannot bend far upwards). The vertebral column is extremely flexible more movable than in representatives

of the preceding families [Canidae, Ursidae; see vol.

II, pt.

la].

The

skull is relatively small, with shortened facial parts

and

broadened, usually swollen braincase. In the majority, the facial portion is strongly shortened and the disproportion of the skull is
noticeable; in some,
is
it

is

more proportional, but

the facial portion

493

always shorter than the braincase. In small forms, the tubercles, crests, etc. of the skull are usually weakly developed, and in large forms, they are very well-defined. The line of the upper profile is
usually straight, without noticeable frontal elevation and the skull
is

low, as

if flattened.

In several, the skull

line.

is

quite high, with the

upper profile an arciform

small, as are the paroccipitals.

The mastoidal processes are usually The auditory bullae are quite large, The
orbit
is

but flattened or only weakly inflated, usually without a septum.

The alisphenoid canal

ral

is

absent.

large,

and the tempo-

fossa

is

of great size.

736

The second upper molar

mula
all
is
I

is

absent.

The complete dental

is

for-

C-P M = 38.

14

This formula

not encountered in

species of the family (developed in martens, Maries; wolverine,

s.s.,

Gulo; badgers

Meles, Helictis, Arctonyx and others).

A
1

re-

duction in tooth number often occurs in the premolars and molars.

This

occurs

in

various

combinations
3
1

- M - = 36
3

(otters, Lutra);

3 = 34 P-M
1

(polecats, weasels, Mustela,

Eyra, Galictis; American badgers, Taxidea; skunks Spilogale,

Mephitis) and even

ilci P M = 32
3
1

(skunks, Conepatus) and

lici
3
1

M - = 32

(ratel,

Mellivora). In one case (sea otter, or

kalan, Enhydra), reduction occurs in the

3

number of

incisors re-

tained,

and the formula takes the form

I-C-P-M- = 32.

13

The

great-

est reduction,

however,
2

is

reached in the number of molars and

1

premolars
3
1

M- = 28

(Lyncodon). In rare cases,

all

or

some of

the premolars are absent, and the dental formula signifi-

cantly varies individually, with age, as well as geographically

dental formula

the

may be

P-P-P-P^4433

= 38 = 36 = 34 (badger, Meles

melesy.

The fourth upper premolar and first lower molar are developed in the form of typical carnassial teeth. The upper carnassial tooth usually has three roots; its inner is well developed. The main outer cusp is not divided. The large cheek teeth, at least along the outer
margin, are sharply pointed (the inner cusps are blunt) or have
blunt tips and tubercules. In the
first case,

the posterior teeth are

'Some contradictions in information on dental formulae of the different genera, which are often found in review works, may in part be explained by this situation.

737
usually elongated with an inner heel and in the second

they are

strongly broadened, irregularly triangular or irregularly quadran-

gular in form.

There

is

curved, divided

an os penis, usually thin and long, sometimes strongly at the end, and without a longitudinal groove. Anal

glands are well developed, sometimes very strongly so, and always

have a sharp, sometimes stupefying odor, especially in those forms which are able to exude secretions (skunks and, to lesser extent

some badgers). The head is usually small

or of moderate length, ears are

rounded, rarely quite large, but usually small and sometimes strongly

reduced and fusing [with the trunk] (in the aquatic forms). The tail is of moderate length, sometimes long, considerably longer than
the hind foot, and even

more than half the body

length; thin or

broad and muscular, with a wide base; in several, it is very short, only slightly extending beyond the dorsal fur. The eyes are of moderate size; but in some, they are relatively large and bulging. The tip of the nose is bare, the lips slightly movable.

more or

less

The pelage,
494

in relation to different kinds of biological types

and the regions of occurrence, is quite variable. Some northern forms have a dense, soft and silky coat which is very valuable as
fur (martens, sable, sea otter, or kalan). In the majority of aquatic

forms

(otters) the

underfur

is

dense, thin and kinky overlain by

coarse guard hairs. The fur of

some forms

is

coarse and bristly,

with poorly-developed underfur or without

it; its

value

is

very low.

Color

is

to the highest

degree variable, from more or less

unicolored dark brown to yellowish-red, black, white, black- and white, etc. In some cases, coloration is very bright and contrasting

from black, red, yellow, brown etc. Not rarely, there are peculiar patterns on the head in the form of stripes and elongated spots. Cases of coloration, where the venter is a dark (black) color, and the dorsum a light gray or white (ratel, badger, and others) or when stripes and spots are found against the basic black background (skunks), are characteristic for the family. In some,
individual and geographic variation in color
is

considerable.

Sexual dimorphism in color

is

absent, but males are usually

in color is absent

somewhat
or
is

larger than females.

Age dimorphism

expressed only
is

in the first

juvenile pelage. Seasonal dimor-

phism

only manifested in northern forms and

variable. In extreme cases, the

may be

very sharp,
is

with regard to length, density and other fur properties. Fur color

somewhat

brown summer coat may


738
be replaced by the white winter (ermine, weasel). There are one or two molts annually.

There are several pairs of teats (2-4), sometimes 1 (sea otter). Concerning general appearance and body structure, the numerous species of the family are extremely diverse, although they may
be divided into several biological types (see below).

feature

common

to all lies in their relatively short limbs.

characterized by a thin

The majority are and elongated body and long neck; some

have a more compact and quite massive build, sometimes a heavy body; the relatively small or small and narrow head is characteristic. Among mustelids there are small forms with a very long, even serpentine body, and narrow head (no distinction between head and body) the purely terrestrial weasels, ermines and kolonok; some are more rugged and large, but also of the polecat

type; their analogs are connected with water, but less specialized

minks; semiarboreal and rocky, good climbers

excellent

swimmers and
in

divers, the truly

martens and ilka*; amphibious fresh-water

and flexible body. Specialization reaches its marine-dwelling sea otter living on the seashores, which possesses pinniped characteristics (structure of the hind limbs).
otters with very long

extreme degree

and short limbs are usually move at a walk, sometimes as if "crawling" or jumping ("galloping"), with the back arched. The extremely specialized aquatic forms move poorly on land. Another extreme type is represented by the heavy terrestrial badgers and ratel, which exhibit various degrees of adaptation to digging, sometimes high, but in any event the highest in the order.
unable to

These forms with long

to very long trunk

move

at a trot

More

or less "neutral" are the

little

specialized forms (South-Asian

badgers), but the

large animals of

are represented by the relatively low mobility with broad massive trunks (Euroasiatic and American badgers Meles, Taxidea). The skunks (vonyuchki*'^) also constitute a particular category of slow-moving but small representatives of the family. This is connected with their remarkable passive defence (exuding a stinking fluid from the anal glands).

more developed

The wolverine

is

large and massive, but a

good climber.

Russian name

for

North American

fisher,

**Literally, "stinkers"

M. pennanti

Sci. Ed.

Sci. Ed.

739
Difference in the size of the different species

is

very great.

The largest forms

attain a

body length of 90

cm

(badger); 120

(South- American otter, Pteronura brasiliensis) and even 150

495

cm cm

(sea otter). The latter has a weight of up to 40 kg. The smallest form has a body length of about 15 cm, a weight of about 100 g rixosa). The ratio of the or less (weasels of the group pygmaea* size of the largest, by weight [to the smallest] is, therefore, 1:400.

All mustelids live either singly or in families; as an exception

they

may form

small groups having a

burrows. Species leading a sedentary

the family (data on

common, or neighboring, mode of life do not occur in

some American
is

otters are not well defined).

Trophic specialization

quite variable. There are trends to-

ward sharply pronounced energetic carnivores feeding on warmblooded vertebrates (chiefly rodents weasels, polecats, etc.) and strongly restricted ichthyophages (otters). With one or another defined conditions, the food of both groups

may be mixed

with other

animals, and sometimes also plant foods. Carnivorous "collectors"

constitute quite a large group, feeding on small

warm-blooded and

cold-blooded vertebrates and on invertebrates, often digging them

out of the ground (badgers, skunks and others). There are also

carnivores which hunt large animals (wolverine

deer, moose) and

even a form specializing

kalan).

in

feeding on sea-urchins (sea otter, or

As

a rule, they are sedentary, sometimes very firmly at-

tached to a place. They usually construct their personal under-

ground shelters, which sometimes have very complicated structure, and inhabit them for many years; sometimes they occupy foreign burrows. Some northern forms hibernate (common and American badgers, skunk). The number of young differs in various groups from 1-2 and up to 10.

They

are mainly nocturnal, but a portion are crepuscular forms.

is,

Of
and

the sense organs, hearing

in

apparently, the most developed,

level of psychological

some, smell

also.

The general

[men-

tal] activity is,

apparently, lower than in species of the wolf and

bear families. In particular, with few exceptions (otters), they are

not easily domesticated and trained. They are definitely found in
all

landscapes

from tundras

to deserts, in

humid

tropical forests,

marshes, rivers and sea coasts. In a vertical direction, they are

distributed

from depressions below sea

pygmea

level to

extreme heights.

*Spelled

in

Russian original

Sci. Ed.

740
Their geographical distribution
entire world.
is

very extensive

nearly

the

The range includes

all

the entirety of the South

Ameri-

can continent, Central and

of North America, including the

whole Arctic archipelago. In the western hemisphere, representa-

on the Falkland and Galapagos on all islands of the West Indies and on some islands in the Bering Sea (Saint Lawrence, Hall, Saint Matthew, Nunivak); the Aleutian and Pribilof islands are included in the range. In
tives of the family are absent only

islands,

Greenland, the range area occupies only the very northern part of
the island east of

Ross

Strait

and

its

eastern coastal zone southlat.

ward, somewhat more southerly than latitude 70" N.

(below

Scoresby Sound). In the Old World, the range includes the whole of Africa, Europe and the mainland of Asia. Species of the family

on Iceland and the majority of the Arctic Islands Novaya Zemlya, Severnaya Zemlya and Wrangel Island), but are encountered in Kolguev, Vaigach and the New Siberian archipelago. In the Far East, the range encompasses the Commander, Karaginsk and Kuril islands, Sakhalin, Japan, Taiwan and Hainan. In southern Asia, Ceylon [Sri Lanka], Sumatra, Bangka (absent on Belitung), Java (absent on Bali and farther east), Kalimantan and Palawan are included in the range. In the entire remaining insular region between Asia and Australia,
are absent

(Spitsbergen, Franz Josef Land,

species of the family are absent^ as well as in Australia itself

and the islands of the southern part of the Pacific Ocean. They are also absent in Madagascar, where the Viverridae are very richly
developed.
In connection with their direct persecution (as valuable fur-

bearing species) or exclusion from cultivated regions, the ranges

497

of

some forms were

the family as a

quite strongly changed. However, the range of whole has not changed substantially in the last

century.
In spite of
its

species richness and especially

its

diversity of
distinctly

forms, the family represents a well separated group.

It is

Canoidea, and

demarcated from other families within the order and in the group its independence does not admit of any doubt. On the whole, these are predators of quite highly, often very highly
specialized type.
^Information from several authors (Weber, 1928; Hilzhelmer, 1930 and others, and even Pocock, 1941) on the distribution of the family eastward to the Philippines (otters), is apparently erroneous (Carter, Hill and Tate, 1946).


742
In their

extreme forms they are only inferior to the cats

the

most definitively predatory of the mammals. On the whole, in their group Canoidea, they are analogous to the viverrids among the group Feloidea, and are in general not inferior to them in their degree of specialization, and in some respects (adaptation to aquatic
life),

considerably superior to them.

genus
cialized.

Nectogale

Among

the viverrids, only one

is

analogous to otters though not more spesimilarity of

The external

some forms of

the family with

individual contemporary viverrids represents only convergence

systematically, both families are quite strongly divergent and be-

long to different groups (see characteristics of the order). Within

the scope of contemporary carnivores, the marten family
related to raccoons (Procyonidae) and
is
is

closely

relatively far

from the

bear and wolf families.

The family
the

itself represents

a quite ancient

group

its

first

representatives were found in the early Oligocene together with

most primitive representatives of the wolf, viverrid and cat They exhibit the greatest closeness to primitive forms of the wolf family. However, differentiation of the mentioned famifamilies.

was, in the lower Oligocene, still not completely determined, and views concerning the most ancient and primitive forms of the marten family are contradictory (Stenoplesictinae are on one hand
lies

assigned to Mustelidae, on the other, to Viverridae). Oligocene

representatives of the family were small animals, true predators

with sharp teeth, apparently, like the polecat or marten. Differentiation of the

main existing branches

is

already occurring in the

upper Oligocene (otters), middle (badgers) and upper (skunks) Miocene and middle Pliocene (wolverines).

The development of

the family proceeded, apparently, mainly

in the northern hemisphere

in

North America and Eurasia the

in

Mustelidae were known from the lower Oligocene, in Africa they

are found

from the middle Pliocene, and

South America

only
is

in the Pleistocene.

Concerning internal structure, the contemporary family

really satisfactory division of
it

ex-

tremely varied, and until now, there has been no accepted and
into subfamilies or in general,

suprageneric groups. Even long ago, division into two subfamilies

was held by many authors (Mustelinae and Lutrinae otters on one hand, and all others on the other), which division is still accepted by many; into 5 subfamilies (see below) and even into 15 (Pocock,

743
1922).

The

latter is, naturally,

unacceptable, but shows well the

that the

diversity of forms, the diversity of characters and complexity of

radiation in the family.

The view has even been expressed

present Mustelidae does not comprise a phylogenetically united

group, but
account;
families)

is

of polyphyletic nature.

The heterogeneous character

if fossil

of the group becomes even clearer

if
is

forms are taken into

the extreme subdivision of existing forms (15 sub-

followed, not less than 15 extinct subfamilies must

also be accepted (Simpson, 1945).

498

most natural division of the family two subfamilies separation of otters, Lutrinae, which are very completely isolated from remaining mustelids. However, at present, until there is a full analysis of the question, it is practically more suitable to divide the family into 6 the extinct Leptarctinae and 5 contemporary ones. It subfamilies is usually considered that there are 76 genera, of which 47 are extinct and 29 existing. This constitutes 38% of the total number of genera in the order and 22% of the existing genera. The actual number of existing genera is, however, somewhat less (see below). The subfamily of true weasels, Mustelinae, includes the following genera^: Mustela (polecats, ermines, etc. Eurasia, North and South America; 15-16 species); Martes (martens, including kharza [yellow-throated marten] Charronia Eurasia; North America; 6-8 species); Vormela (marbled polecats Eurasia; 1
It is

most probable,

that the

will turn out to be into

species); Eira (tayra

(grison, including
cies);

North and South America; Grisonella North and South

species); Galictis

America; 2 spe-

Lyncodon (South America; 1 species); Ictonyx (African polecat Africa; 2-3 species); Poecilictis (Africa; 1 species); Poecilogale (Africa; 1 species); Gulo (wolverine Eurasia and

North America;

species).

The subfamily of honey-badgers, Mellivorinae, includes one

genus Mellivora (Africa, Asia;
1

species).

The subfamily of badgers, Melinae, comprises the following Eurasia; 1 species); Arctonyx genera: Meles (Old World badgers
(hog badger

eastern
list

Tibet and southeastern Asia;

is

species);

'The following
transferring

of genera

not in accord with data of Simpson (1945). His

revision based on various data led to reduction in the

number of genera, mainly by

some

to the rank of subgenus.

It is

apparently possible that further reduc-

tion will take place.

nized by Simpson,

Genus Grammogale Cabrera (Cabrera, 1940, 1957), not recoghere set aside. The number of species established from different sources, cannot be considered precise. Actually, they are probably somewhat fewer.
is

744
Taxidea (American badger
4 species).

North

America;

species); Melo-

gale, including Helictis ("polecat" [ferret] badgers

southern Asia;

The subfamily of skunks, or vonyuchek, Mephitinae, includes

the genera: Mephitis (North America; 2 species); Spilogale (North America; 2-3 species) and Conepatus (North and South America;
7 species).

The subfamily of
era:

otters, Lutrinae, includes the

following gen-

Lutra (including Lutrogale, true otters

(giant otters and South America; 10 species); "clawless" Amblonyx; (including Aonyx species); America; 1
otters

Eurasia, African, North South Pteronura

Ocean;
1

Africa,

Asia; 2 species); Paraonyx (Africa; 3 species);

Enhydra (sea

otter, or

kalan

northern Pacific
the

species).

In the entire family, therefore, there are

24 genera and about 70

It is

species. This constitutes about 1/3 of the species of the order.

more probable that there are about 65 number of genera and species, the family

species.
is

As regards the richest one among

the group Canoidea, and one of the richest in the order. It is only inferior to the viverrids (about 36 genera and 75 species) and greatly exceeds the number of the species of wolves and cats. Of the total number of genera of the family, four are African (not counting Mustela, which penetrates to the northwestern edge); North American, two South six are Asian and Eurasian, three

American (Grammogale is not taken into account); 2 Afro-Asian, 3 North and South American; 1 distributed in Eurasia, North in Eurasia, the Americas and Africa. and South America; and 1 Therefore, as regards the number of genera, the fauna of the Old World and especially of Asia is the richest and most heterogeneous. As regards the number of species, the poorest is shown to be the fauna of North America (5 endemic species and several com-

mon to the other continents), and the richest endemic species and several others common

Eurasia (about
to the

17

America and genera, endemic only two Africa). It is remarkable, that in spite of both monotypic, the fauna of South America is relatively rich in
species (about 16), mainly on account the diversity of otter species

and skunks of the genus Conepatus*

The
vores.

practical significance of the family

if

is

very great.

It is

one

of the most important,

not the most important, group of carni-

The species of

the family play a particular great role as

*In Russian original, misspelled Canepatus

Sci.

Ed.


745
fur-bearers. If the chinchilla

is

not counted, the family comprises

the most valuable fur-bearing animals of the existing fur species

martens, fisher, otters, and most of

all,

sable and sea otter (kalan).

Some
499 ing

of them have even played a great historical role

the open-

up of Russian Siberia was first of all connected with sable, whereas the northern Far East and the shore of the northern part with sea otter. The of the Pacific Ocean, including the American penetration of Europeans into northern North America from the south, was also determined chiefly by furs, among which an important role was played by American sable [marten], fisher and

mink. Exploitation of mustelids played an important role in the life of natives of northern Eurasia and America, and the importance of
sable in ancient Russia and to present times
is

generally known.

Representatives of the family play one of the chief roles in the

economics of the fur trade at the present time. Beside the particularly valuable species mentioned, there are a considerable number of other less expensive species, but which are
obtained in large quantities such as polecats, ermine, Siberian weasel, mink, wolverine, pama*, various otters, etc., which give
very large amounts of fur every year. Otters themselves represent

main owing to
the
ers, etc.)

fur species in tropical

their natural conditions, are very

and equatorial countries which, poor in fur-bearing

animals. There are

some

less valuable fur-bearing species (badg-

which, in
at the

total, also

play a certain role. Finally, the very

great,

and

present time, the main product of fur farming

consists of

color forms (mutations).

American mink raised in various, artificially obtained The mink is now a more important furis

producing animal than the silver-black fox. Sable, martens, and skunks
are also raised in captivity, but their importance

very small.

Many
sels,

species specializing in feeding

on small rodents (wea-

ermine, polecats, and others) are themselves considered im-

portant regulators of agricultural pests, and of reservoirs and carriers

of dangerous infections (plague and others). Otters play a

role in their capacity to exterminate fish

known
to fish-

and cause harm

farming. Finally, a few species serve as objects of sport hunting.

The

fat

of

some

is

used

in folk

medicine.
are represented 4 out of 5 sub-

In the fauna of the

USSR
23%)

families (80%), 8 of 24 genera

(33%) and 17 species (not counting

(V.H.).

acclimatized) of 70 (about
*Meaning unclear

Sci. Ed.

746

Key for
1

Identification of

Genera of

the

Family Mustelidae

(2).

Digits of the forelimb conjoined, and entire

sents a single unit.

paw

repre-

gated wide flippers, covered with fur dorsally and ventrally. In each half of

Hind limbs transformed into elonin which digit V is longest, and evenly

lower jaw, 2 incisors. Upper surface of molars and premolars smooth, with rounded edges, without cutting
tubercles or sharp cusps

the tubercles are strongly blunted

genus of sea
2(1).

otter or kalan,

Enhydra (page

1330).

Digits of the fore and hind limbs are not conjoined and

movable (sometimes perhaps connected by membrane),

hind not forming flippers and middle digit
[III] is

longer

than others. In lower jaw, 3 incisors on each side, molars

and posterior premolars with well developed pointed

bercles or with sharp cusps.
3 (4).

tu-

External ear conch absent or rudimentary; entire lower

body, limbs, side of the head and muzzle black, upper

side of
fields

body

is

grayish-white, color boundaries of both

sharp. Molars and premolars 4 in lower jaw.

The

last tooth

of upper jaw considerably smaller than

preceding'* (Figure 185)

genus of honey-badgers, Mellivora (page 1207).

500

(3).

External ear conch present, color of one sort or another.

Molars and premolars 5-6

5 (6).
light, dirty-gray

in

lower jaw.

Ventral surface of body black or blackish-brown, dorsal

(without yellow color). Posterior upper molar 2-3 times larger than preceding one. Body length more than 60 cm genus of badgers, Meles (page 1228).
6
(5).

Color different,
sal,

if

ventral color black or darker than dor-

then back has a clearly developed yellow or yellow-

ish tone. Posterior

upper molar not larger than preceding

one.

(8).

Digits united by broad

pletely devoid of hairs,

membrane which

is

nearly

com-

on hind foot extending

to claws.

Soles of fore and hind feet bare below. Tail long (not less
than half length of trunk with head), base broad and

muscular. Dimensions large

body

length about 70-75

in

cm. Last molar tooth approximately equal

^Crown viewed from above.

dimensions

747

Fig.

185. Characteristic structure of posterior part of toothrow in upper

jaw of some

genera of the weasel family, Mustelidae. Sketch by N.N. Kondakov:

1

Genus

of honey-badger, Mellivora; 2
otters, Lutra;

Genus of badger, Meles; Genus Genus of wolverine, Gulo.

3

of

to

preceding one. First upper premolar pushed away from

lies

toothrow and
8 (7).

at inner side

of canine (Figure 186)

otters,

genus of

Lutra (page 1283).

Swimming membranes between

end of

digits absent, or are very

weakly developed, covered with hairs and do not reach digits. Food pads*, at least those between digits, with hair. First upper premolar lies posterior to canine. Tail of different lengths, but not muscular and base not
broadened.
9 (10). Dimensions are large

body

length more than 70 cm,

condylobasal length of skull more than 130

relatively short (about
501

mm.

Tail

20 cm). Coloration evenly brown with light bracket-form bands of different intensity

*Misspelled

in

Russian original

Sci. Ed.

748

Fig. 186. Position

of canine and

first

genus of

otters, Lutra.

premolar of the upper jaw of representative of Sketch by N.N. Kondakov.

passing along sides and across croup from shoulder to

shoulder
10(9).

Dimensions smaller

genus of wolverines, Gulo (page 920). body length less than 65 cm,

condylobasal length of skull less than

variable; coloration otherwise.
11 (12).

15

mm.

Tail length

In upper

jaw behind canine

molar), in lower
side of

5 teeth (4 premolars

and

(4 premolars, 2 molars).

On

posterior

the lower

main cusp of last premolar (third tooth from rear) jaw has a small additional cusplet (Fig. 187) genus of martens, Maries (page 749).

Fig.

187.

Last lower premolar tooth of genus of martens, Martes (sable, Martes

zibellina L.) with additional cusplet

on main cusp. Sketch by N.N. Kondakov.

12(11).

In

upper jaw behind canine, 4 teeth,

in

lower

5.

Additional cusplet on posterior side of last premolar (third

tooth from behind) of the lower

jaw

absent.

13(14). Back

is

parti-colored

and

stripes,

yellow with irregular dark spots or dark, densely covered with irregular light

spots and stripes. Posterior processes of pterygoids reach

tympanic bullae and united with them (Fig. 188) genus of marbled polecat, Vormela (page 1176).
14(13). Coloration otherwise, without spots or stripes. Processes
of pterygoids do not reach tympanic bullae genus of ermines and polecats, Mustela (page 954) (V.H.).

749

Fig. 188.

Position of pterygoid processes in genus Vormela (joined with tympanic

bullae) and in the other genera of the weasel family (not joined with tympanic

bullae

black

[common]

polecat, Mustela putorius L.). Sketch

by N.N. Kondakov.

502

Subfamily of Weasels
Subfamilia Mustelinae
Gill,

1872

Genus of Martens

Genus Martes
\115. Martes. Frisch.

Pinel, 1792
d.

Natur-Syst.

vierfuss.

Thiere, p.

11.

According

to decision of the International

in this edition

Commission on

Nomenclature, names
significance.

have no nomenclatural
1, p.

1792. Martes. Pinel. Actes Soc. Hist. Nat. Paris,

domestica Pinel = Martes foina Erxleben.

1829. Zibellina.
1,

55. Martes

p.p. 31, 34.

Entw. Gesch. u. Natur. Syst. Europ. Thierw., Mustela zibellina Linnaeus.

p.

1865. Charronia. Gray. Proc. Zool. Soc. London,

flavigula Boddaert.

108. Mustela

1928. Lamprogale. Ognev.

Memuary

Zool. otd. Obshch. lyubit.

estestvozn., 2, p. 26. Substitute for Charronia Gray, 1865,

preoccupied by Charonia Gistel, 1848. (Mollusca). (V.H.).

Species of moderate dimensions.
Skull relatively narrow and elongated, without sharp crests and

protuberances. Braincase relatively quite large. Facial portion relatively elongated,

laterally interorbital

zygomatic arches weak, not strongly diverging, and postorbital constrictions weakly defined.


750
Supraorbital processes small.
slightly convex.
infraorbital

Upper profile of skull gently sloping, Convexity of fronto-nasal region at level of foramena weakly developed. Diameter of infraorbital
to

foramena almost equal

diameter of alveolus of canine. Bony

auditory bullae are quite large and their inner parts noticeably bulging towards one another. Hamate processes of pterygoid bones
not united with auditory bullae. Mastoid processes small, weakly protruding laterally, lateral occipitals (paroccipitals) well

developed.

Bony

palate wide.

Dental formula
First

14 1--3
1

= 38.

lus or its traces remains.

premolars very small and sometimes shed, but an alveoOn main apex of last lower premolar

(anterior to carnassial tooth

third tooth posteriorly), there

is

small additional cusp on the inner side.

On

inner side of median

lower carnassial tooth (first molar second tooth posteriorly) there is no additional cusplet. Well developed pointed apex on inner blade of upper carnassial tooth (P4). Trunk is elongated, but not to an extreme degree, or modergeneral body structure fairly slender and relatively ately short

apex

of

proportional. Tail quite long

in various species

from

1/3 to 2/3 of

body
broad

length.

Head comparatively

large with large protruding ears,

almost triangular in form, facial part pointed. Large ears and sharp muzzle give head a short, broad wedge-shaped form characteristic of "marten" appearance. Limbs digitigrades
at base,

and hind feet broad. Fur dense, long, soft and silky (one of the most valuable furs). Tail covered with long hairs, fluffy. Color, in majority, is unicolor brown tones; in one case it is bright and patchy, with a combination of white, black, yellow and brown. On the throat and chest,
fore-

there
503

is

usually a lighter area.

Seasonal dimorphism in character of fur very pronounced; in winter, soles of the fore- and hind feet are entirely or almost entirely

covered with dense

hairs.

Seasonal variations in color

slight.

Sexual differences
erable;

in color are absent;

but dimensions of males

average larger than females, and in several forms are quite consid-

sometimes (M. pennanti) sexual differences exist in fur characters (fur of females is softer and finer and, consequently,

more expensive than

that of males).

751

There are two pairs of inguinal teats (subgenus Martes). All martens are very active, quick, flexible and clever animals.
All are adapted not only to a terrestrial, but also to an arboreal

mode of life, some to a very complete degree (able to descend head first down a trunk, their hind feet capable of being turned backwards, etc.). Because of their relatively short limbs, they move on land by jumping, with their back arched. They are forest and
montane-forest animals, also inhabiting unforested mountains;

however, they do not

live in

extremely high mountains (nival zone).

They

are strictly sedentary, and

8.

monogamous.

Litters

from 1-2 and

are predators,

3-4, up to

Shelters are mainly in hollows and in tree trunks, but

also in rock clefts.

They

are solitary animals.

They

feeding on flesh, chiefly of small rodents, but sometimes also on

warm-blooded animals, even small ungulates (yellow-throated They also utilized lower vertebrates and invertebrates, and in the ration of several, a significant role is played by plant food (berries, nuts). Torpor or hibernation does not occur.
larger

marten).

In dimensions, the species of this genus are quite homotypical.

The majority have a body length of about 40-50 cm and a weight of up to 1800 gm. Two species are large the yellow-throated marten, M. flavigula, has a body length of about 75-80 cm and a weight more than 2.5 kg, and the fisher, M. pennanti a body

length of about 70

and The range of the genus

cm

a
is

weight up

to

6 kg.

vast and covers a considerable part

of the temperate and cold-temperate zones of the northern hemisphere, part of Central Asia and the region of subtropical and
tropical forests of southeastern Asia.
It

occupies

all

of Europe, the

greater part of Asia, and the northern half of North America. In

Europe and Asia the range extends from the northern limit of forests [southward] to Spain, Italy, Greece, Crete, Asia Minor, Syria, Palestine, Iran, West Pakistan, Kashmir, Punjab, Himalayas, Indochina, the Moluccas, Sumatra, Banka [Island], Java and Kalimantan
(Borneo).

small part of the range, in southern India,

(for details, see

is

cut off

from the general area of habitation

descriptions of individual species).

below under

In North America, the range in the north

is

limited by the
is

northern border of forest vegetation. The southern border

repre-

sented by a meandering line, extending from the Atlantic coast

ginia, northern

westwards through the states of Connecticut, New Jersey, VirNorth Carolina and Tennessee, Indiana, northern

752
Illinois,

northeastern Iowa and Minnesota (except the southwest),

northeastern North Dakota, the southern part of Saskatchewan and the southeastern corner of Alberta. The range occupies all of Brit-

Colombia and from there and from Alberta, the range gives off two large extensions southward along montane regions. One exish

tends along the coastal regions

states of

along

the western parts of the

Washington, Oregon and to the central parts of California, and the other, more deeply into the region, extending along the eastern borders of the states of Washington and Oregon to Idaho, the western parts of Wyoming into Utah, Colorado and to

Mexico. In the range is included Newfoundland, Cape Breton Island, Vancouver, the Queen Charlotte islands and others along the Pacific coast of America northward to 60 N. lat. (absent on Kodiak and the Aleutian islands). In sketching the boundaries in the steppe and desert regions of Asia, conthe northern parts of

New

siderable gaps exist in relation to natural conditions; in Siberia, the

range

is

considerably changed in some places due to

human

activ-

ity; there are blank areas

and the northern border does not corre-

505

spond everywhere to the restored range cited. Within the limits of the family, the genus itself constitutes a well-isolated group, characterized by a series of features, most of

by the dental formula. Craniologically, the genus is entirely uniform and the species within this category of features are characterized basically in details. The genus is well delimited from closely related genera, in particular Mustela. The uniting of these genera is a
all,

former matter.

It is

an entirely whole group, not only

in relation to

its morphology, but also in an ecological and zoogeographical sense. At present, the independence of this genus is not doubted. Moreover, attempts to divide the true martens, united here in the genus Martes, into a separate subfamily Martinae, which were done in the past, and which made sense only in the case of extreme subdivision of the family, cannot be considered established, and

are

genus of true martens (our martens and the American, and sable), Martes, and that of the yellow-throated marten, or Himalayan marten, Charronia. According to a series of characteristics the yellow-throated marten is sufficiently well differentiated from the true martens; however,
it

now rejected. Many authors

divided the genus into two

the

is

more

correct to separate

it

only as subgenus. There are no

essential craniological differences in the yellow-throated marten.

g
kS<C

g
.'

__^
_^_^^j

^^
/

1)

'

/*

^^.^

^St | ^ \^
^^;wiiO

-"-^

5^j3
\
1

\
<D

^%~^

.S

h^
)

ii^/

'^
1

>
-1

1
1

"".

'

\:^.
S
^^


754
Therefore, in the genus are two subgenera

Charronia with spe-

cies and Martes, comprising all the remaining. The determination of the position of this genus within the
limits of the family, given the diversity of
difficult.
its

radiation,

is

quite

This group

is,

in

some

respects, highly specialized.

How-

ever, with respect to the relative completeness of the dental sys-

tem,

it

may be

conditionally placed at the beginning of the series

of genera of the family, and of the subfamily Mustelinae.

Concerning

its

origin,

Martes belongs

to a

number of primitive

genera of the subfamily; fossil representatives of the genus are

known from the lower Pliocene (according to some data from the Miocene) of Europe and Asia. The closeness of the connections between the primitive forms are not clear starting from the lower Oligocene, a quite large number of geiiera of the subfamily are known, mainly from the Miocene. In particular, the genus Mustela, to which the genus Martes is quite close, is known from the upper Miocene. The number of the species in this genus is not yet fully estab-

M. zibellina; Japanese M. melampus; pine marten, M. martes; American marten, M. americana; stone marten, M. foina; ilka (fisher-marten or pekan), M. pennanti; kharza [yellow-throated marten], M. flavigula and South Indian kharza, M. gwatkinsii. The actual number of the species is, apparently, less, not more than 6. M. melampus is only a subspecies of M. zibellina, and M. gwatkinsii is only a race of M. flavigula. M. americana is closely related to the sable and to the pine marten, which, in their turn, are very closely related to each other. The relationship between these species requires further clarification. The 6 species of the genus constitute about 8.5% of the species of the family and 17-20% of
lished; usually eight are admitted^: sable,

sable,

the species of the subfamily.

The

greatest

number of species
fisher,

are

endemic

to Eurasia (4). In

America, there are two: the

marten, M. americana.

M. pennanti and the American

America,
this

Both

in Eurasia, as well as in
it

genus has great

practical significance, since in

are included very valuable fur-

bearing species

all

true martens

and among them, the sable. The

in this

importance of the yellow-throated marten

great.

respect

is

not

The kharza plays some

in the

role as a destroyer of
is

game

animals,

^Reference to 10 species

genus (Novikov, 1956),

evidently based on the

value of such "split" species, as in Miller (1912).


755

and

in

Europe the martens

the pine and

in particular the stone

cause harm to the hunting economy and to poultry enterprises. In the USSR are found 4 species: 3 of the subgenus Martes:
sable M. zibellina

1)

Linnaeus, 1758; 2) pine marten, M. martes 506 Linnaeus, 1758; 3) stone marten, M. foina Erxleben, 1777, and

one of the subgenus Charronia: 4) yellow-throated marten, M.

flavigula Boddaert, 1785.

They

constitute

66%

of the species of

the genus and about 1.2% of the fauna of the country. Species of this genus are distributed over the entire forest zone of the Union and in the mountains of the Caucasus, Middle Asia

and southern Siberia.

They

are valuable fur-bearing species (V.H.).

Key for
1

Identification of Species in the

Genus Martens

(2).

Color patchy there are portions of white, brownish-yellow, blackish-brown and bright golden-yellow colors. Tail length constitutes about 2/3 of body length. Dimensions
large

condylobasal
may
exist,

length of skull of adults

more than
(p.

100
2(1).

mm (body length

of adults to 75-80 cm)

yellow-throated marten,

M.

(Ch.) flavigula

905).

Coloration uniform
spot

reddish-brown or brownish;

a light

or bright-yellow color, or head

on throat and chest only, white, yellowish may be lighter than body,

sometimes even whitish. Tail length comprises less than condylobasal 2/3 of body length. Dimensions small length of skull not more than 90 mm, adult body length
not more than 60 cm.
3 (4).

Tail relatively short.

Its

length with terminal hairs less

it

than half the body length or equal to

it

hardly extends

beyond the ends of the extended hind limbs. Light throat spot absent, or not clear, small and has irregular form and not sharply demarcated from is not precisely outlined color of neighboring parts of body. Top of head usually

lighter than back.

Bony

auditory bullae elongated and

at

adjacent. Distance

between them

middle of their lengths

less or equal to half of distance

from anterior point of

zibellina (p. 757)

bulla to posterior margin of lateral occipital process

sable,

M. (M.)

4(3).

Tail relatively long. Its length with terminal hairs

more

than half the body length

it

extends beyond the ends of

756
the extended hind limbs for

more than

1/4 of

its

length.

Throat spot variable

sharp outline.

in

form, but well-defined and with

color as that of back.

Top of head same

Auditory bullae shorter and widely separated. Distance between them at middle of their length more than
half of distance from anterior point of bulla and posterior

5 (6).

margin of lateral occipital process. Throat spot color pure white and usually gives off two posterior projections extending to the forelimb. Inner part
of upper molar
is

507

not wider or only slightly wider than

190). Tapering,

the outer (Figure

posteriorly pointing

projection along posterior border of sphenopalatine notch

absent, or
it

is

barely noticeable. In winter pelage, a bare,

spotted pad

is

observed among the hairs on sole, and somestone marten,

times digital pads^

M. (M.)foina

(p.

874).

Fig. 190. Last premolar (carnassial) tooth and first molar of upper jaw of sable (left), Martes (Martes) zibellina L., pine marten, Martes {Martes) martes L., and stone marten, Martes (Martes) foina Erxl. Sketch by N.N. Kondakov.

6 (5).* Throat spot varies from light-yellow to orange, rarely

white; usually spot gives off posterior projection between forelimbs.

Inner part of upper molar considerably wider than

outer.
ally

Well-marked posteriorly pointing projection usufound on posterior border of sphenopalatine notch. In winter pelage the digital pads on soles hidden in hairs forest marten, M. (M.) martes (p. 825) (V.H.).
form and color of throat
spot,

'Differentiation between stone and pine martens by

though most frequently practiced,

essential.
*

is

not always reliable (see section below "Descrip-

tion" of species). For purposes of identification, characteristics of teeth structure are

In

Russian original, "5(5)"

Sci. Ed.

757

Subgenus of True Martens

Subgenus Martes
Pinel,

1792

SABLE
Martes (.Martes) zibellina Linnaeus, 1758
1758.

Mustela zibellina. Linnaeus. Syst. Nat. Ed. X. L p. 46. Northern Siberia. According to Ognev (1931) and other

1840.

more precisely: western Siberia north of Tobol'sk. Mustela melampus. Wagner. Schreb. Saugeth. Suppl., 2,
authors,
p.

229. Japan.

1844.
1855^.

Mustela brachyura Temmink. Siebolds Fauna Japon. Mamm., p. 33. Japan, Hokkaido. Mustela zibellina var. asiatica. Brandt. Mem. mathem., phys. et natur., 7, p. 6, 23, Taf. 1, Fig. 1. Kamchatka.^
M. zibellina
5.

1855.

var. alba. Brandt.

Ibidem,

p. 14,

Taf. 2. Fig.

1855. 1855.

M. zibellina

war. fusco-flavescens. Brandt.

Ibidem,

p. 14,

Taf. 2, Fig. 6, Taf. 3, Fig. 7.

1855.

M. zibellina var. ochracea seu ferruginea. Brandt. Ibidem, p. 14. Taf. 3, Fig. 7. M. zibellina var. maculata. Brandt. Ibidem, p. 14, Taf. 3,
Fig. 9.

1855. 1855.

Mustela zibellina
Taf. 2, Fig. 2.

var. asiatica rupestris. Brandt.

Ibidem,

Mustela

zibellina

var.

asiatica

sylvestris.

Brandt.

Ibidem, Taf.
1918.

2, Fig. 3.

"^Mustela zibellina subsp. kamtschadalica. Birula. Otchet

deyatel'nosti Ross. Ak. Nauk, p. 82. Kamchatka.

collection of works reprinted from volume VII of ^Date for individual volume "Memoirs" (see list of references at end of book). The pagination is given according to the same edition. 4n the work of Brandt mentioned, the individual or "ecological" (mountain, forest) trends of the sable are described. By the name, "var. asiatica", Brandt means all Asiatic sables in contrast to American sables ("var. americana"). At the same time,
the
in

Table

1,

Fig.

is

illustrated a

normally colored sable, originating, as stated

in the

legend from Kamchatka. Formally, under such conditions, the

clear that Brandt under his var. asiatica

name
this

asiatica

may be
it

ascribed to Kamchatka sables. However, since in the whole sense of the paper

is

had

all

Asian sables

cannot be done.

*In Russian original, not in chronological order

Sci. Ed.

758
1922.

508

1922,

1922.
1922. 1923. 1925.

Mustela zibellina princeps Birula. Byalynitskii-Birula. Ezheg. Zoologich. muzeya. Ross. ak. nauk 1917-1921, 22, p. 8. Barguzin range. Mustela zibellina var. kamtschatica. Dybowski. Arch. Tow. Nauk. Lwowe, 1, p. 349, Nomen nudum. Kamchatka. Mustela zibellina var. baicalensis. Dybowski. Ibidem, p.
349.

Nomen nudum.

Vic. Baikal.

Mustela
349,

zibellina var.

amurensis. Dybowski. Ibidem,

Mori. Joum.

p.

Amur, Ussuri. Mustela melampus coreensis*. Kuroda

mal. 4,
p. 27.

et

Mam-

Tenan, South[em]. Chusei district]. Korea.

Martes zibellina yeniseensis. Ognev. Journ. Mamm. 6, No. 4, p. 277. Krasnoyarsk u[yesd], Yenisei guber[naya],
taiga plain.

1925.

1925. 1927. 1927. 1941.

Martes zibellina sajanensis. Ognev. Ibidem, p. 278. Orzybai [=Orsyba] river, northern Sayan [mountains]. Martes zibellina sahalinensis. Ognev. Ibidem, p. 279.
Sakhalin, Vedernikovo. [=Saghalien, Wedernikovo]. Martes zibellina correensis. Kishida. Choju Chosahokoku,
4, p.

130. Korea.

Martes zibellina hamgyensis. Kishida. Dobuts Zasshi, 39,

p.

509.**

Martes zibellina tungussensis. Kusnetzov. Tr. Mosk. zootekhnich. inst., I, p. 116. Lower Tunguskaya [river].
M.iartes) z.(ibellina) sahalinensis arsenjevi. Kusnetzov.

1941.

Ibidem,
1941.
Ibidem,

p.

122.

Samarga River, Ussuri

Territory.
.

M.{artes) z-iibellina) sahalinensis schantaricus

p. 122. Bolshoi [Great] Shantar Island,

Kusnetzov.

Okhotsk Sea.

1943.

Bazhanov. Bashanov. averini. zibellina ^Martes in SSR Souza 1942, Kazakhskii fil. Ak. N. p. 13. Katon-Karagaiskii region, Vostochno-Kazakhst. obl[ast],
southern Altai.

1947.

Martes zibellina altaica. Jurgenson. Yurgenson. Tr. Pechoro-Ilychskovo*** Zapov. 5, p. 179. Oirotskaya
" auton. obl[ast], Altai.

*In Russian original, misspelled koreensis

**No

type locality cited in Russian original

in

Sci. Ed. Sci. Ed.

'Released apparently,

1944.

***In Russian original, misspelled "Ylychskovo" Sci. Ed. '"Type (Zoological museum, Moscow university) from left bank of middle course
of Bystrukha river, upper Katun [river]; southwestern Altai (V.H.).

759
1955.

Martes zibellina tomensis. Timofejev et Nadejev. Timofeev et Nadeev. SoboF, p. 37. Kuznetskii Alatau.
Tutuyas river

tributary of Tom

[river].

1955.

1955.

Martes zibellina angarensis. Timofejev et Nadejev. Timofeev et Nadeev. Ibidem., p. 41. Angara valley in Boguchaev region, Krasnoyarskii territory. Martes zibellina ilimpiensis. Timofejev et Nadejev. Timofeev et Nadeev. Ibidem., p. 44. Kochechumo watershed tributary of Kotuya [river], 67 N. lat. (about 100"

E. long.).

1955.

Martes zibellina vitimensis. Timofejev et Nadejev. Timofeev et Nadeev. Ibidem., 47. Valley of the Mama
river, right tributary of the Vitim [river], Yakutiya. Martes zibellina obscura. Timofejev et Nadejev. Timofeev et Nadeev. Ibidem., p. 47. Upper Chikoi river right tributary of Selenga [river], Trans-Baikaliya.

1955.

1956.

"M.(artes) M.(artes) z.(ibellina) jakutensis. Kusnetzov

(1941)". Novikov Khishchnye mlekopitayushchie fauny

SSSR,
509

p.

185. Yakutiya,

Aldan Valley" (V.H.).

Diagnosis
Tail length with terminal hairs not

more than half the body

length.

Color uniform, on throat and chest a yellow spot of irregular form;

sometimes absent. Head usually lighter than back, sometimes

whitish.

Bony

auditory bullae relatively elongated and adjacent.

is

Inner half of upper molar

wider than outer (V.H.).

Description

The
It

sable, in its winter fur,

is

a very graceful

and elegant animal.

has a moderately elongated body and relatively short limbs with

"In the referenced book by G.A. Novikov [1956; see Lit. Cit.], the "Yakut sable" was designated as that set forth in quotation marks. Nevertheless, in the work of B.A. Kuznetsov (1941) to which Novikov refers, the Yakut sable was not given a name, and was designated as "7) Martes zibellina s. sp.?. Yakut sable" (page 120). A short preliminary diagnosis was given by B.A. Kuznetsov, and it was shown that the "Yakut sable" lived mainly in the valleys of the Aldan and Kurchum rivers". Since B.A. Kuznetsov did not give a scientific name to the Yakut form, its description must be formally considered that which has been given by G.A. Novikov and must be named M. (M.) zibellina jakutensis Novikov, 1956. As for type locality, since "Yakutya" (mentioned by Novikov) is too broad and ill-defined, it is hereby restricted to the Aldan valley.

760
very broad, thickly furred
feet.

The

sole pads of the digits and feet

tail is

and the claws are hidden

tively short,
its

in the

dense hairs. The

compara-

length usually constituting about 1/3 of the body

its

length and not exceeding half of

length. In the "skinned"*

its

animal
slightly

it

is

small (no

more than
is

1/4 of

length) and extends

It is

beyond

the posteriorly extended hind legs.

densely

covered with long hairs and

very fluffy. The head of the sable

appears very large, cuneate in form

the nasal
tip.
is

part tapering, but

slightly extended; the ears are large, upright, of triangular

form

with a broad base and a somewhat blunt

The eyes

are bulging,

quite large, black; the naked tip of the nose

are black.

black and the claws

Since the trunk of the sable

is

elongated, and the legs are

its

relatively short, the animal always arches

hind parts are often higher than the shoulder; the

back strongly, its tail is always

very well-

fluffy. Nevertheless, the sable in its winter garb, is a

proportioned animal, an appearance which has nothing in

the sable confirms

common

with a small cat. The general impression from the appearance of

its

extraordinary activity and speed, deftness

and confidence of movement. The sable runs quickly, moving by jumps ("trotting"; it almost never walks, due to the above-noted proportions of its body), it can make great jumps, can climb trees
and
cliffs

well and can dexterously

make

its

way among
in.

the stones,
it

fallen trees, etc. Finally,

due

to its broad, densely haired feet,

can confidently traverse loose snow without sinking

load of the sable track
is

The weight

less than in all

our other species of the

entirely differtail.

genus.

The appearance of
ent

the sable in

summer garb

is

it

looks thin and long, with longer legs and a thinner

The

head seems disproportionately large, with huge ears, and the neck

and long. The relatively great breadth of the feet in the sable draws attention to it. The summer pelage of the sable is short, coarse and sparse, and the underfur is weakly developed. The length of the guard hairs on the rump (croup) is 20-30 mm with a thickness of 100 to 105 microns; length of underfur is 12-14 mm with a thickness of 16-18 microns. For each guard hair there are 5-6 underfur hairs;
is

thin

summer

the
to

number of
600-700.

hairs in a

cm^ area of skin on the rump

is

equal

Literally, "in flesh"

Sci. Ed.

761

The winter pelage of

hairs

the sable

is

dense, with a large quantity

of underfur, exceptionally fluffy and silky. The length of guard

on the croup equals 36-48 mm, the length of underfur, 2428 mm. The thickness of the guard hairs is 85-90 microns, of

underfur, 14 microns. Therefore, winter underfur, with a length

almost double that of the summer, and are even absolutely thinner; such proportions are also true for the guard hairs. For one guard
hair, there are
1

20-25 underfur

hairs; the total

is,

number of

hairs in

cm^ area of skin on the rump

tail

on the average, 13,500

(Kuznetsov, 1951, Yenisei sables; Pavlova, 1951). In winter pelage

the hairs of the
are considerably longer, denser and fluffier

(they reach 85

mm)

and the

feet are also

covered with denser and

longer hairs. In

summer pelage

the claws and pads of the digits

and soles can be seen, but in winter they are completely hidden. The winter sable walks on a furry cushion. The great luxuriance of
the fur on the feet in winter considerably increases the area of the
feet,

510

and facilitates the animal's movement on loose snow. While the winter fur of the sable is entirely variable in color

(see below), the

summer pelage

of the animals in a single locality

is

and

in various parts

of the range,

uniform. Even in very light

races (for example, Tobolsk sable), the

Its

summer

fur

is

very dark.

monotone: dark-brown, darker along the back, slightly lighter on the sides, and still a bit lighter on the belly. The tail is blackish-brown. The legs have the same tone as the sides or
color
is

they are somewhat darker than the color of the back.

On

the throat

and along the lower neck, there is completely undefined outlines, spreading onto the sides of the neck. In some, it is absent. Between the ears and eyes, a dark area extends to the nose; the cheeks, the region in front of the ears, the
ears,

a lighter yellowish area with

and behind the ears

is

is

occupied by a light area of ochreous

color merging posteriorly with the lateral parts of the throat spot.

This pattern

head
hairs.

is

sometimes absent or weakly defined, and the whole darker. The ear margins are trimmed with short ochreous
the

The generally dark color of

so

summer

sable

is

explained not

much by

the coloration of the various hair categories as by the

structure of the fur.

The underfurs have

ratio of

a yellowish-brown color,

the guard hairs and contour hairs are dark-brown or almost black.

The previously mentioned

both hair categories (5-6

1)

762
thus results in the general color of the fur being mainly determined by the color of the contour hairs (Pavlova, 1951). The winter pelage is characterized by a lighter color of the underfur bluish-gray at the base and sandy or brownish at the tips. The guard hair has the same color as in summer, but with a predominance of black pigmentation over the brown; however, the number of the down hairs for each guard hair is much greater (see above), and the general color, to a great extent, is determined by them (Pavlova, 1951). The deviations are more strongly expressed

in the general

tone of the color of the underfur. In this respect, in

one and the same place, individual variation in color has a much greater range in winter than in summer, and geographic variation in color is considerable, and much greater than in summer. The winter fur of the sable has a quite uniform color over the entire body. The sides and the lower surface of the body has a

somewhat
tail

less intense color, but the contrast

is

insignificant.
is

The

has the same color as the back, but the terminal half

usually

darker.

The

legs,

and especially the

feet, are

darker than the back.

The

anterior part of the head and ears or the

whole head

is

conthe

siderably lighter than the remaining parts of the trunk.

throat and neck there
is

On

a large light area, yellowish in color,

without sharp outlines.

With

this

type of color distribution,

its

general color

is

sub-

jected to very great variation bearing both an individual and a

geographical character. The general color tone within the species

varies

from sandy-yellow

to brownish-black.

The character of

the

throat patch also varies as well as head color and the degree of

uniformity of color of the different parts of the body. Sables having either very

monotone

color, or

more contrasting colors occur.

described above
is

The darker color of back and

istic

tail

characterits

of light-colored sables

the darker the

is it

sable, the less

conthe

trast,

and

in the darkest sables, not noticeable.

So also with

color of the head

in the majority of cases,

it is

light-grayish and
as the back; the

only in particularly dark sables

the

same color

cheeks are always slightly lighter. Sometimes, even in relatively dark sables, the head is light, occasionally almost pure white. The
throat patch
is

very variable. In

many

animals,

it

is

completely
all

absent or only represented by a tiny, weakly colored part of the

fur.

Sometimes, on the contrary, the throat patch with

is

its is

variation in shape,

sharply outlined.

The color of

the patch


763
511

usually yellowish, but there occur light-gray and even white. This
or the other tendency in the development of the patch
carry a geographical character.

may

also

A
In
it
it,

century of practice in the Russian fur business has elabo-

rated a very precise categorization of color variation in sable fur.

no attention

is

assigned to the quality of the fur

all

itself,

and

is

applied and adapted to

the geographical forms (sorts) of

sable.

These

sorts are as follows:

Golovka. Color particularly dark. All fur pitch-black or blackish-brown. Guard hairs black with barely noticeable brownish tinge.

Underfur dark-bluish without

light hair tips.

Throat patch weakly

defined or has form of small orange "star".

skin color pitch-black with

Head

dark.

Skins of golovka are divided into vysokaya golovka (extra)

almost pure black guard hairs;

normalnaya golovka
black guard hairs.

skin

color blackish-brown with brownish-

Podgolovka. Color

lighter.

Spine and sides of skin dark-brown

or dark-chestnut color without well-defined reddish tinges on sides.

Guard

hairs

dark-brown or dark-chestnut. Underfur gray (blue)

tips.

with chestnut hair

Head

grayish, lighter than spine. Throat

patch ill-defined, not bright.

Skins of podgolovka are divided into vysokaya podgolovka

skin color dark-brown with dark-brown guard hairs and bluish-

gray underfur, the hair

tips

of which have dark-chestnut

tints;

and

normalnaya podgolovka ish guard hairs and bluish-gray underfur with chestnut hair tips. Vorotovyi. Color medium-dark. Skin surface dark-brownish or moderately intense brownish with a dark stripe along the spine and lighter, slightly reddish sides. Guard hairs on spine dark-chestnut, sides are lighter. Underfur grayish with reddish-brownish or dark
sandy-yellow hair
bright.
tips.

skin color dark chestnut with dark brown-

Head

light,

grayish. Throat patch large,

vorotovyi

Skins of vorotovyi sable are divided into temnyi [dark] skin color dark-brownish with weakly defined reddish

tinges on sides, guard hairs chestnut, underfur with reddish-brown-

and normalnyi [normal] vorotovyi skin color brownon the sides, guard hairs light-brownish, and underfur with sandy-yellow hair tips. Mekhovoi. Color light (light-brownish, sandy-yellow, or pale-yellow). Guard hairs brownish or light-brownish. Underfur
ish hair tips;
ish with clear reddish tinges

764
light-gray with reddish or yellowish hair tips.
gray. Throat patch ill-defined, large.

Head

light, grizzled

With regard
sable
is

to all
at

remaining qualities, the skin of the mekhovoi

about 15-20%, the dark vorotovyi

evaluated

35%, and

the vysokaya

podgolovka

at

30-

at

60-65%

of the value of the

vysokaya golovka skin (Kuznetsov, 1952). Golovka and podgolovka categories comprise the group of "dark" sables, vorotovyi is designated as "medium", and mekhovoi

as "light". The scheme given obviously characterizes the correlation of color of different parts of the body, particularly the color

tone of the head.

Amplitude of individual variation

races

in different

populations and
is

may

differ. In several

parameters the range of variations

par'^

ticularly great,

and the difference between the extreme forms

is great.

512

This or another percentage of relationship between basic color

types in different parts of the species range, with other equal conditions,

well characterizes individual geographical races and

populations (see below, section on "Geographic Variation"). In

several races, separate types
sables, there are

may

rarely be found. Thus, in Tobol'sk

no golovka, and in some Trans-Baikal, light sables are rare, and golovka constitutes up to 70%. At the same time, the numerical ratio of color forms within one race may change from year to year, and sometimes this change is fairly considerable. The multiyear average is, however, maintained. In Tofalar region of Irkutsk district, these fluctuations had the following form

(Nadeev and Timofeev, 1955) (Table 48).

Table 48. Change in the ratio of color forms within one race
Winter
Total

Skin quality

number

Dark (%)
16.4
19.3
8.8

Medium (%)
67.6

Light (%)
16.0
14.8

1941/42 1943/44 1944/45 1948/49 1949/50

1950/51

269 130 295 327 711

901

65.9 67.6 63.0

64.7 76.9

23.6
8.0 5.8

29.0
29.5
11.3

11.8

511

'^At the International fur auction in

Leningrad

in

1961 the most expensive skin

of the extremely valuable Barguzin sable was sold for four hundred and five dollars,

and the very cheapest of the same race, for twenty dollars (B.A. Kuznetsov)
of 1:20.
It

a ratio

is

natural that, in evaluation, all the characteristics of the skin played a

signal role (fluffiness, softness, delicacy, luster of the fur, grizzling, size etc.).
ever, color
is

How-

one of the main characteristics.

765
Similar to this type of change in fur color of various races and
populations, there are, apparently, also changes which take place
in

one direction for a quite extended periods. Thus,

in regions of

Siberia adjacent to the Yenisei beginning at the end of the 1940's

the color of the sables gradually started to lighten, and
at the start
still

continues

of the sixties. Even the release of dark Barguzin sables in some regions did not prevent this process (Tashtinsk, Yeniseisk and Turukhansk regions, 1949-1957). In some parts of the range, a darkening of color in the population is observed (K.D. Numerov).

There may occur

in skins

an admixture of pure white guard

hairs, giving the beautiful "grizzled" fur. This

admixture

may be
absent

great ("grizzled sable")

or

negligible,

or completely

("glukhaya sable"). White sables (albinos), chromistic and skewbald (with white spots)
countered.

may be found
tint as

as exotypic variations.

Sables with a bright orange

well as gray

may

also be en-

Sexual and age difference in the color of sables

is

absent.

Young sables in their The skull of sable

facial

first
is

autumn

are clad in adult coats.

relatively

narrow and elongated. Both the

and particularly the braincase are elongated. The distance from a median line joining the ends of the postorbital processes to the alveoli of the middle incisors constitutes more than 65% (but less than 80%) of the distance from the same line to the posterior point of the occipital crest. The muzzle is relatively narrow. Nasal bones have usually fairly sharp constrictions in their middle portion.

Zygomatic arches are

relatively thin, the supraorbital proc-

esses are well developed and broad, and the postorbital constriction

wide (more than the width of the skull above the canines). The mastoid processes do not protrude beyond the lower margin of the auditory meatus. The sagittal crest is only developed on the posterior-most part of the cranium and the occipital crest is weakly developed. Bones of the auditory bullae are relatively swollen in a longitudinal direction and close to each other: the distance between them in their median point of their length is less (rarely equal to) than half the distance from the anterior end of the chamber to the posterior edge of the paroccipital process. The auditory
is

tubes are well developed.

514

The longitudinal diameter of

In this latter the inner blade
is

the upper carnassial tooth

is

approximately equal to the transverse diameter of the upper molar.

considerably larger than the outer.

766

513

Fig. 191. Skull of the sable,

Maries (Martes) zibellina L.

767
Sexual differences
variation
in the skull are insignificant

and represented

only in the somewhat smaller dimensions of the female skull.

is

Age

quite great. Besides the increase in general dimen-

sions, with age the skull acquires a more elongated form, and the zygomatic width increases as does interorbital width. On the other

hand, the postorbital constriction of the skull (behind the supraorbital processes), narrows.

The

interorbital

width

in

males 8-9
is

months old constitutes 18.5

older than 3 years
18.0 and 15.5
is

mm

on the average, while an animal

20.2, corresponding postorbital constriction

mm. With

age the so-called "temporal lines" which

outline the region of attachment of the masseteric musculature to the braincase, gradually approach each other, and the sagittal crest
is

formed on the
According

skull

by the time of sexual development of the

and other characters,
192;

animal.
to the character of these lines

the following age groups of sable are distinguished (Fig.

Nadeev and Timofeev, 1955).

Fig. 192. Schematic outlines of sable skulls of different ages. From Nadeev and Timofeev, 1955. A) Age group I 8-10 months; B) Age group II 1 year, 8-10 months; C) Age group III 2 years, 8-10 months; D) Age group IV about 3

years,

8-10 months and

older.

,.-,

768
i.e. middle of first and extend parallel to each other, only sometimes approaching each other at occipital crest, but do not fuse. Form of skull is rounded. Occipital crest is rarely visible, and the sagittal is absent. Incisors not worn, or very
I

Age

young animals 8
life.

to 10

months

old,

winter of

The temporal

lines separated

slightly obliterated. Ratio of interorbital

width to postorbital in

males, 0.84-1.17

Age
winter of

II

about

(M
1

1.06); in females,

0.78-1.14

(M

1.06).

life).

8-10 months old (middle of second Occipital crest well developed. There is a rudimenyear,

tary sagittal crest, temporal lines

come

together posteriorly form-

ing an acute angle. Incisors slightly worn. Ratio of interorbital

width to postorbital
1.24

in

males, 1.04-1.22

(M

1.10); in females, 1.04-

(M Age

1.11).
III

about 2
come

years,

8-10 months old (middle of

is

third win-

ter

of

life).

Occipital crest

moderately or strongly developed.

Temporal

lines

together in blunt angle at anterior half of

braincase, in posterior part form well-developed sagittal crest. Ratio

of interorbital width to postorbital in males, 1.09-1.37

in

(M

1.20);

females, 1.08-1.33

(M

1.17).

Age IV about 3 years, 8-10 months old (fourth winter of life and older). Occipital crest strongly developed, sagittal crest is moderately or strongly developed it extends along the entire brain-

case; temporal lines absent (evident only in interorbital region).

Ratio of interorbital width to postorbital in males, 1.14-1.55

1.30); in females, 1.17-1.43
515

(M
all,

(M

1.24).

The majority of

the

above-mentioned features, and above

the position of the "temporal lines" and the

development of

crests,

are determined by increase in mass of masticatory musculature with

aging, and

by

their

other. If in the

growth upward from both sides to meet each young animal (Age II) this musculature is relatively
left sides

small and muscles of the right and

are separated, then by

Age

III, they are closely appressed along the sagittal plane and lie on the braincase as a solid elongated mound (Fig. 193). The tail skeleton consists of 15-18 vertebrae. The os penis is

at its distal end, and the ends of this fork may be directed toward each other, forming a half-ring, but they do not close the

forked

ring.

The length of

the ossicle in Trans-Ural sables

is is

39.2-M

41.4-43.2

mm

(Yurgenson 1947)'^ Intestine length

in the structure

1000-M

"Other features

of this bone given by

Ognev (1931) and used by

Novikov (1956)

are not constant and not characteristic.

769

Fig.

193. Position of masticatary musculature in different age groups of sable.

[Fig.

Groups [A, B, C] are those given on in previous figure and Timofeev, 1955.

192] from

Nadeev

1790-2700 cm, which constitutes 250.0-M448-630.0% of the (365-M422-490 mm). Heart weight is 5.23-M8. 82-15.20 gm, or 9.0-Ml 2.2-2 1.5%o (n 117); lung weight (without trachea) 8. 10-M 14.9-52.50 gm; liver weight 10.32-M14.90-45.7 gm; kidney weight 1.40-M2.76-5.35 gm; spleen weight 0.62-M 1.65-3.90 gm'^ (Timofeev and Nadeev, 1955). The diploid number of chromosomes 38 (V.N. Orlov). Dimensions (extreme variants for the species) of sable are as follows: length of body of males, 375-580 mm; of females, 320510 mm; tail length of males, 110-170 mm; of females, 90-176 mm; hind foot length of males, 70-105; of females, 60-90 mm; ear length of males, 50-56, of females, 43-55 mm.
length of the carcass without the hide

Condylobasal length of skull of males, 74.1-94.6 mm; of females, 70.0-84.4 mm; zygomatic width of males, 38.7-56.2 mm;
of females, 32.9-52.8

mm;

skull height of

males (together with

auditory bullae), 27.0-37.4

material of

mm;

of females, 26.5-39.6

mm

(after

Nadeev and Timofeev, 1955). Weight [in gm] of males (22) of Yenisei sables in January, 1110-M1472-1810, of females (21), 820-M 1132-1560. Weight in July in males, 1200-M1433-1810; in females, 870-M980-1300
(Kuznetsov, 1941). Optimal weight of breeding sables before the
'The relative weight of

all

organs

is

overestimated, since
Its

it

was determined
is

according to the weight of the carcass without the hide.

[hide] weight (n 230)

440-1300

gm (M

730.5).

770
beginning of rut (15 July)
in captivity

(Pushkin fur farm): males

with body length between 39 and 49 cm, 1400-1900 gm; females with body length between 36 and 46 cm, 900-1400
1947).
516

The maximum known weight of

male

is

1869

gm (Starkov, gm (V.H.).

Systematic Position

The sable

is

a typical representative of the genus Martes, closely

all

related to the true martens; as with

of them,

it is

quite sharply time, the

distinguished from the fisher, M. pennanti. At the

same

characters of the forms zibellina, martes, foina and americana are

combined in individual species in quite complex assortments which do not permit arranging them in a successional series. However, it is evident that of our two martens, the pine marten (M. martes) is more closely related to the sable. This is indicated not only by a
series of similar features in their morphological characteristics, but

also by the hybridization of these species in nature Ckidas"^^), and by the fertility of the hybrids in the back-crossing. Notwith-

standing

all

of that, obviously the species independence of sable

in

and marten are accepted

It is

what follows.
it

possible that the American marten M. americana belongs

to the species zibellina, although

has several marten features

(bones of the auditary bullae). In

any event, the sable, pine marten

and American sable themselves represent a close group. It is quite possible that the sable should be considered the less specialized form of this group and thus (perhaps together with the American sable) the least specialized species of the genus (V.H.).

Geographic Distribution
Forest regions of
Siberia, northern Europe,

Mongolian Republic,

northeastern China, the Korean Peninsula and Japan.

Geographic Range

in the Soviet

Union
its

This constitutes the overwhelming part of

great part of the state territory
the northern

range.

It

occupies a

all

the forested part of Siberia

and

European

part of the

USSR.

'^For details about kidas, see the section on pine marten.

771

Determination of the natural range of sable presents great

ficulties. In the

dif-

was subjected to intensive pursuit over the past thousand years, and in Siberia, over the past few hundred years. By the twentieth century not only had the number of sable decreased a hundred-fold, but also the general extent of its range was very sharply reduced. The sable is associated with forests, first of all with taiga; however, it is ecologically quite flexible and within the forest zones is met with under entirely different conditions from it thickets of prostrate nut-pines in the high mountains to low land swampy taiga, and from forests of the Amur type to forest-steppe islands and riparian forests. This allows one to think that, in the past, the sable occupied the entire forest zone of Siberia. There are
the sable

European part of the

USSR

indications of

its

occurrence in the forest-tundra,

at least the

south-

ern extreme, not only in the past, but also at the present time.

One may

consider that the natural range of the sable was con-

tinuous. There were, of course,

more

or less significant openings

and gaps in it, related to natural landscape conditions (for example, Minusinsk steppe). However, they themselves constituted exceptions, and their area was immeasurably smaller than regions inhabited by the sable. Reduction in the range of the sable was quite unique and took place in two ways. On one hand, mainly in the south, but also in the west and the northeast there was recession of the limits of the
range.

On

the other hand, within the boundaries of the range, great

in

expanses were formed

which the sable has completely disapwas the chief way the range was reduced the general areal extent of the sable's disappearance is no larger an area than that in which it
peared. Thus, the range was broken into separate parts. This

persists.

The separate areas inhabited by

into "islands", usually small,
cially in the east, they are so

the sable

became transformed

and

in the majority of cases, espe-

completely isolated from each other

that their natural restoration, given the cost of restocking,

517

seemed

impossible. Along with their direct pursuit and destruction, a large,

although not so significant, role in the reduction and fragmentation

of the sable
is

range was played by the felling, burning and plough-

ing up of the taiga, which

scale,

was gradually taking place on a large

and

in particular forest fires.

772

The
century,

picture of the range described at the beginning of the 20th

was

the result only of destructive

human
is
is

activity.

As

in-

dicated by experiences in our day, the sable

quite viable and

extinct, as

completely "contemporary" and, by no means, some have attempted to portray it.

going to become

The destruction of sable

scarcity of historical data

started so long ago, that, with the

all,

above

about Siberia

to restore the

picture of

its

range

is

now

quite difficult. Proceeding

from the
territories

ecological characteristics of the sable, in regard to

some

one may consider that the actual range in those places was greater than that drawn on the basis of contemporary data and historical
information.
It is

particularly difficult to establish a picture of the

range of the sable in the European part of the

USSR. The views

presented by different investigators about the previous limits

of sable distribution in this part of the country are entirely at
variance.

After reaching

its

minimum and extreme

of fragmentation at

the beginning of the first decade of the 20th century, the range the beginning of the second decade, during from 1912 on of complete prohibition of its hunting and in connection with other circumstances. By the third decade, it was again reduced. By the end of the 30's and in the 40's and 50's, not only did the number of sable increase by several times, but its range area had strongly extended as a result of legal protection, conservation and reintroductions. Protected areas inhabited by sable have increased, uniting several previously isolated areas again, and new sables appeared in those places where they had been long absent. In this way, in the last ten years the sable's range was found to have changed rapidly. This also makes the definition of

increased

somewhat by

the period

distributional data quite difficult.

The
tor

history of the range of the sable

is

a clear

example of a
its

range change occurring under the influence of the anthropic fac-

at first, its

deep, uncontrolled destruction, and then

planned

restoration. Naturally, the range of sable cannot

to

be fully restored

its previous size, because in some places natural conditions have been so intensively changed that they became unsuitable for the existence of the species. However, at the present time, not all possibilities have been employed to restore the previous range of

the sable.

773

The
there
is

outlines of the westernmost part of the range in the past

(historical time)

cannot

now be

reliably

and completely restored.

If

more

or less reliable information, not disputing the exist-

ence of disagreements, along the Urals and the northeastern European part of the Union, westward to Severnaya Dvina then the
question of the occurrence of sable farther to the west
clear. Its discussion dates
is

not so

back almost one hundred years; never-

theless, in quite distant times as well as

more recent

years, very

competent authors (Middendorf, 1867; Sabaneev, 1875; Zhitkov 1937), regarded information on the occurrence of sable in the European north, northwest and west with great scepticism or denial.

In recent years, however,

new

materials have been found

(Kirikov, 1952, 1958, 1960) which reopened

anew
is

the discussion

of this question'^
519

The

localities farthest

west for which there

mention of the

presence of sable are located in Lithuania, Byelorussia and Smolensk oblast. Besides Lithuania in general, the sable was also noted in Vil'nyus and Kaunas, Zelenaya Forest in the former Ponevezhsk county and near Knyshin city (now in Poland to the
northwest of Belostok
first

the westernmost

point; in the 16th

and the

half of the 17th century). In Byelorussia, Minsk, Lutsk

district in the

(now

Volynsk
N.

Ukraine; the southernmost point, about 51

lat.), Novogrudok (Middendorf, 1867), Polotsk, Vitebsk, and Grodno were mentioned as places inhabited by sable in the 18th century; in the 1880's, sable was noted in Belitsk and Klimovich

counties; in the latter, at the border with Roslavl' county of

Smolensk governance. There is mention of the occurrence of sable around Smolensk at the beginning of the 17th century, and in 1668, in Bryansk forest, which was contiguous to the south with
Byelorussia. Therefore, old data on sable occurrence in Lithuania and Byelorussia are substantively reinforced. In the north, to the west of the Northern Dvina, sable were

noted within the limits of our country on the Sun' river, which
'*Here data

on

the occurrence of sable in the west are laid out very briefly and

schematically, chiefly after Kirikov, 1952, 1958, 1960, as well as Middendorf, 1867,

Zhitkov, 1937;

Polyakov, 1873, Sabaneev, 1875; Pleske, 1887; Ognev, 1931; Yurgenson, 1933, Van den Brink, 1958 and several other sources. Separate references are only given in the case of their absence in S.V. Kirikov. It must be taken into
consideration that the data of Sabaneev (1875) on westward transgressions of sable,

which are commonly used (Ognev, 1931 and others) are taken from Middendorf (1867) and contain little that is original.

774

518

Fig.

194.

zibellina L. in Europe. V.G. Heptner:

Data on the past and present distribution of the sable, Martes (Martes) 1 generalized mentions of concrete, and 2

the occurrence of sable during the

16th and

17th centuries; 3

generalized concrete, and 6 generalized information for the 18th century; 5 probable northern and southern limits of the information for the 19th century; 7

range in the northern European part of the country in the past; 8

concrete,
limits

and

in

the

(contd.)

775
flows into Onezhsk [Onega] lake from the northwest (middle of
the 17th century)'\ in Lapland (in the 17th and 18th centuries),

and particularly on the Kola [Peninsula]

century, where in 1834
it

at the

end of the 17th

found somewhere in the northern parts of present Karelia (Kemsk county of Arkhangel'sk governance), and in the former Kholmogorsk county, and in the 16th century, in Velikii Ustyug district. This is the southernmost place
still

was

of occurrence of sable in the section of the northern European part of the

USSR

under consideration.
is

Concerning Finmark, the sable

northern

mentioned

in

Scandinavian

folklore in the 9th-10th centuries. Finally, northern Finland and

Sweden hosted

the occurrence of sable in the past (about

300 years ago). Here, the range forms an irregular triangle lying at the eastern border of Finland, approximately between 65" N. lat. and Lake Inari; its apex is located somewhere a little south of Kirun in Sweden (Van den Brink, 1958). The materials given above show that the range of the sable in
and Kola Peninsula, and extended to Finland and Sweden. Concerning the districts lying to the east of the Dvina and Ural [rivers], the Cis-Urals and the Urals [mts.] themselves, there is a body of sufficiently accurate information indicating a considerably wider distribution of sable in the past. In the 16th-17th centuries,
the
the west occupied the taiga of Arkhangel'sk district, Kareliya,

the sable was, apparently, widely distributed along the

its

Mezen' and

western tributary, the Vashka (Udor),

i.e.

evidently also in the

and Dvina. Concerning the Mezen' and Vashka, there is also information up to the middle of the 19th century. In the north, sable extended to Pustozersk in the lowermost Pechora, and approximately along this latitude the northern border
interfluve
it

between

of

its

range reached to the mouth of the Ob'

river.

In the

Vychegda

basin, sable existed and

were

still

quite

com-

mon
(Fig.

at the

beginning of the 17th century, being found not only

194 contd.)

extreme northwest in the past (Van den Brink, 1958); 9 probable southern limit of the range in the Neolithic (Yurgenson, 1933 with modifications); 10 recent (in 1960) western limit of sable range. In Byelorussia and Lithuania, in Kareliya and on Kola

Peninsula,

all

data points in literature are presented, in remaining sections

individual peripheral points.

only
said:
...

'
"would
it

the manuscript of the

monk

Epifanii

minor companion of Protopop

Avvakum whose

personal report of the place

if
...

is

ascribed to one hunter,

who

be any wonder,

God

sent precious sable or fox or other animals

all

animals and birds to live here" (report of A.N. Robinson).

776
along the right (Yarenga,
(southern) tributaries

Vym' and

others), but also along

its left

the Sysola as well as Bolshaya and Malaya

even farther to the west.

Vizinna*, Lop'yu and others (Kirikov, 1958, 1960). Even in the

19th century, sable lived in the Vishera basin, in the neighborhood

of Cherdyn' and in Cherdyn' county, along the Chusovaya, and in

earlier times, they lived at these latitudes

Sable also occupied the western parts of present

Komi ASSR

and all of former Permsk governance, i.e., the territory to the east of 54 E. long., which nearly corresponds to present Permsk district (Sabaneev, 1875). Approximately to this line or a little to the west, there are mentions referring to Biserovo at the sources of
the

Omutninsk, on the Ponino* a little north of Glazov, at the sources of the Vyatka and in the region of Sarapul (Shaberdin and Strel'tsov, 1930; Yurgenson, 1933). The supposition concerning the occurrence of sable in the eastern half
to the northeast of

Kama

520 of the

former Vyatsk governance (Sabaneev, 1875) is supported by documented places of occurrence; however, concerning more western regions, i.e. to the west of approximately the 47-48 meridian [E. long.], concrete data are absent. (Data of Shaberdin and
Strel'tsov are, however, not very definitive, since they report the

capture of kiduses even in 1928/29).

Farther to the east, the border was deflected, turning south and proceeded into Krasnoufimsk, embracing the forests along the Tyusha, B[olshaya] and M[alenkaya] Sarsa, B[olshaya] Sarana and Kashanka [rivers]. Still further south, there is evidence on the occurrence of sable in the 18th century in the Urals in the upper

Ufa

river (Pallas, 1786)

Urals to

and along the southwestern slope of the the east of the city of Ufa even in the first half of the 19th
true southern border of sable distribution in the Urals in

century (Sabaneev, 1875)'^

The

the time under review, has apparently, extended

somewhere

in the

*Not found

Sci. Ed.

'^References sometimes occur in literature on the occurrence of sable in the past

near the city of Ufa. This

original sources. Pallas (1786) writes: "It

secondhand rewritings and simplification of sometime happens, though quite rarely, that along the Ural mountains, and especially in Kama, and along the upper Ufa river outliers of sable appear (page 25; information dated from 1770). Sabaneev (1875, page 201) reported that the last sable, killed "on the southwestern slope of the Urals", was near Berezovka village, Ufimsk county.
is

the result of

777
region of 55-56 N.
(Pallas, 1786)
lat. the sources and upper reaches of the Ufa and Kaslinsk mountain (Sabaneev, 1875)^^. In summarizing everything said, it is possible to accept that, in
still

historical time, and,

in part at the

end of the 18th and even the

19th centuries, the southern boundary of the range of sable in the

European
lat.,

part of the

USSR
in its

crossed the Ufa

began in the Urals at about 55-56 N. upper or middle course, extended to the

the mouth of the Belaya, passed thither to Vyatka and farther to the region of Velikii Ustyug. Thence, it turned directly westwards to Olonetsk territory (southern Kareliya), passing somewhere through the region of Lake Onezhsk [Onega], possibly through its middle, or even southern, part^. the upper

Kama somewhat above

Beyond
noting,

this

line,

it

is

possible

that

separate
It

areas
is

of

occurrence were found in Lithuania and Byelorussia.

worth

however, the complete absence of information from Moscow, Ryazan and other middle-Russian regions, about which there are many historical documents, and in general, about the whole extensive area between Lithuania and Byelorussia, on one hand, and Arkhangel'sk and the Olonetsk forest, on the other. It is
possible that in previous centuries, a separate focus of sable inhabitation, isolated (artificially?)

from the main range was found

in the
re-

southwest, similar to several of the present Siberian outliers.

ality is that
it

The

lies to the

south of the southern border of Neolithic sable

fossil finds as

the

known to us at the present time (see below). The northern border of the range extended, apparently, along northern border of the forest in the west, Kola was mentioned
they are
in

and

the east
its

Pustozersk,
i.e.

and along the border of the forest

lying to

east,

north of the Arctic circle. In the Urals, the

northernmost point inhabited by sable was, apparently, the Voikor

river

an

Ob' tributary

at its

mouth on

the Arctic circle^'.

"Sabaneev's assumption (1875, page 200) that it extended to 52 "perhaps not based on factual material. Rychkov (1762) directly affirms that "sable occurs nowhere in Orenburg province (which at that time included the South Ural; V.H.)." To
N.
lat.", is

postulate the existence of sable south of 55-56,

is

now

only possible by analogy to some

authors after Sabaneev,

other northern species which descend along the Urals south as far as 52, and for very

remote times. Evidence concerning

on misunderstanding,

5,

accepted by

many

is

based

this parallel

passes south of the Ural river along the steppe.

^"There are data on the occurrence of sable around Arsk northeast of Kazan in
the 16th century (the manuscript of Prince Kurbskii; after Kirikov,

1960), but this

point remains strongly isolated spatially at present.

^'All places, especially

where no reference was

cited, are taken

from S.V. Kirikov

(1952, 1958, 1960). The boundaries herein, however, have a different outline than in
the

map

of S.V. Kirikov (1960).

778
521

The interpretation suggested here of the previous range of sable Europe agrees well with the limits of the range in the Neolithic, which generally corresponds to the outline of the [present] southin

ern limit of the sable's range, but considerably

more

to the south.

This line (Yurgenson, 1933 with modifications) passes from the

northern part of the Gulf of Riga across the lower Velikaya river,

above Pskov, proceeds a little south of Lake Il'men' and somewhat more to the north of Seliger and extends through the Kimp region. Thence, it descends to the southeast, embracing Meshchera, proceeds somewhat more to the north of Kasimov, crosses the Oka [river] above Murom, the Sura in its lower reaches and the Volga
above the mouth of the Kama. This recent boundary crosses in its lower reaches and again the Vatka at about 50 E. long. crossing the Kama above the mouth of the Belaya, reaches the
slightly

Urals.

The substantive point

in the line described is that

it

passes, in

the west, approximately intermediate to the described northern

points of possible habitation in the Lithuanian-Byelorussian region

and the southern limit of the region of previous inhabitation in the area between the Urals and Scandinavia. Evaluating the previous range of sable in North Europe, it must be emphasized that it is, in a zoogeographical sense, regular. Some Siberian taiga species analogous to sable, such as northern
red-backed and gray red-backed voles (Clethrionomys
rufocanus),
Siberian
in the north rutilus,

C.

wood lemming (Myopus

schisticolor) and in part

chipmunk (Tamias

sibiricus) extend very far to the west

all, except the chipmunk, to Scandiof our country However, the presumption of sable occurrence in Lithuania and Byelorussia is not predicated on this type of range. All the mentioned materials, even if the evidence concerning Lithuania and Byelorussia is disregarded, demonstrate the fault in the view of both old and new authors that the sable was not dis-

navia.

tributed far to the west in the past^^.

The present

(50' s) western limit of the range of sable takes the

it

following form. In the north,

begins slightly to the north of the

"Middendorf (1867) affirmed that sable never existed to the west of Syktyvkar Nadeev and Timofeev (1955) believed that "in Europe, sable ... in the past penetrated a very small distance". Zhitkov (1937), Poluzadov (1955) and others held the same view. Sabaneev (1875) did not exclude the possibility of far
(Ust'-Sysol'sk).

penetration of sable to the west, but doubted

it

and only Pleske (1886) believed

in the 18th century.

in the

existence of sable on the Kola Peninsula

still

779
sources of the Lyapin at approximately the latitude 63 30' N.
(a little south of the limit of nut-pine)
lat.

and passes south along the western slope of the Urals through the upper Shchugor, Ilych, Pechora (including the Pechoro-Ilych preserve), Un'ya and to the sources of the Yaiva. Thence, the range boundary turns sharply to
the east, passing through the basin of the

Kakva

river a little north

of the city of Serov, crosses the lower Loz'va and continues to the

Tavda and farther, towards the upper Noska river which flows into the Irtysh below Tobol'sk. It is possible that boundary line extends (or not long ago) to include the region to the west where the Usa
flows into the Pechora, and southward, the Troitsk-Pechorsk
gion (V.Ya. Parovshchikov).
re-

To

the north of the sources of the

Lyapin,

it

passed to the southeast in the general direction of

Berezovo on the lower Ob' (Shvarts, Pavlinin and Danilova, 1951; Nadeev and Timofeev, 1955; for more detailed delineation of the
Ural center of sable occurrence, see Fig. 198).

The northern border of

the range in Siberia begins in the Urals,

on the Voikar river near the Arctic Circle and passes eastward across the lowest reaches of the Ob', apparently, approximately
along the tree-line,
i.e.

somewhat south of

the southern shore of

any case, this line included the basin of the right Obsk tributary of the lower Ob', the Kunovat river (65 N. lat.). Along
Bay. In
the Taz, the range extends to the Arctic Circle and, perhaps, a
little

across

it

(V.N. Skalon).

From

the Taz, the border passes to

the northeast, including the basins of the B[olshaya] and M[alaya]

Kheta
522

(left tributaries

where

in the

Yenisei to

of the Yenisei); crosses the Yenisei someDudinka region and rises along the right bank of the the region of Lake Kit (of the Noril'sk group) and upper
It falls

only slightly short of Noril'sk and here at40' 68 N. (Podarevskii, 1935) or even includes tains a latitude of Lake Pyasino and the upper Pyasina at 70 30' N. (Kirikov, 1960).
Noril'sk creek.
Further, the sable is known at Volochanka on the Kheta, i.e. approximately around 70 50' (Orlov, 1930). To the east, on the Khatanga, the border goes, apparently, to about 70 or slightly

more

to the north, crosses the Khatanga approximately at a latitude 69 30' (Nadeev and Timofeev, 1955) or more to the north (Staraya;

A. Romanov, 1941). It reaches the Anabar, approximately at a latitude of 70 50', on the Olenekat 69 30', the Lena at the same
latitude or at 69 (finds are

known

at

Zheldong 250

km below Zhigansk;

Tugarinov, Ivanov and Smimov, 1934) and on the Omoloi.

780

The range boundary crosses

the

Yana somewhere below

Verkhoyansk, Chondon and Khroma (Kirikov, 1960) crosses the Indigirka about 69 or a bit southward (D. Ivanov) and reaches the Kolyma below Sredne-Kolymsk. On the right bank of the Kolyma, the range encompasses not only the Omoloi basin, but also the Bol'shoi and Malyi Anyui (Middendorf, 1867). From the Anyui, the range border crosses the Anadyr' covering, apparently, its upper and middle course, and across the Khatyrka and Opuka basins, reaches the coast of the Bering Sea in the region of the mouth of the Opuka, a little to the south of Cape Navarin (Samorodov, 1939; Portenko,
1941).

probably about 68", includes the basins of the

Siberia

The description given of the northern border of sable range in was constructed on the basis of direct evidence of its conits

temporary distribution and

It is,

distribution in relatively recent past.

from the 16th century; Kirikov, 1958, 1960), that in the expanse from the Ob' to the Kolyma, the actual natural range boundary ("reconstructed")
however, well founded (archival material dated
coincided with the northern forest limit (see map). Therefore, be-

tween the Ob' and the Yenisei, it lay a little to the north, and between the Yenisei and the Kolyma, significantly more to the north than as shown. The described section of the border east of
the

Kolyma corresponds
zone

to the reconstructed

[border]. In

some

places on the northern border, the sable also lives outside the [taiga]
forest

in

krumholz,

in

deciduous forests of river valleys

(Koryatsk land).
constitutes the eastern range

(Anadyr') and

in nut-pine thickets

The shore of

the Pacific

Ocean

boundary. Within the range are Shantar and Sakhalin islands.

the Kuril islands, the sable
is

Of

have resided on Iturup and Voronov, Kunashir (Snou, 1902; 1963), and to the north, on Paramushir (Kuznetsov, 1949). The latter was shown, evidently, to be mistaken (Voronov, 1963), on Karaginsk Island, the sable is
to

shown

absent, as on the

the

Commanders. The southern border of the reconstructed sable range began in Urals in the upper Ufa at a latitude 55-56 (see above) and
border passed through the whole of eastern Siberia in the

entered Siberia near Chelyabinsk (Pallas, 1786). Further to the

east, the

forest-steppe zone.

went along the so-called "Isetsk province", apparently through Kurgan [city], or between it and Yalutorovsk, and farther to the city of Ishim or south of it and
it

From Chelyabinsk,

781

reached the Barabinsk forest- steppe. Here the range boundary went
to the

mouth of

the

Om,

apparently, north of
at

Lake Ghana and

is

reached Lake Ubinskoe and to the Ob'

border in the 17th century, and,
(Pallas, 1786;
in

Novosibirsk. This

the

some

places, the 18th century

From Novosibirsk,

Sabaneev, 1875; Ognev, 1931; Kirikov, 1960). the boundary turned sharply to the south
in the Biisk

and passed towards the foothills of Altai, apparently

region. Here, the border turned back to the west and then to the

south and southeast, embracing the Altai. In this region, the boundary again passed along the edge of the foothill forests at the
lati-

tude of Biisk; farther,

it

passed around the Tigeretsk range to the

524

north, continuing somewhat to the east of Zmeinogorsk, and then from the western end of the range, along the Uba, Bukhtarma and Ul'ba basins, almost reaching the Irtysh. In the south, the boundary included the Kurchum and the region of Lake (Ognov, 1931; Yanushevich and Blagoveshchenskii, 1952; Afanas'ev et. al., 1953; V.G. Heptner). At the bound-

ary of the range reached the state border".

Altai portion of the sable range

,
It is

probable that the

It

was somewhat broader.

However, there
is

may be

that sable lived also in the ribbons of forest in the Cis-Altai steppes

and extended south on this point.

In the

to the Irtysh.

no information

expanse from the southern Altai to the Pacific Ocean,

the sable range reaches the state border (Tannu-Ola, for instance)

or
as

beyond it. In Ussuri Territory, the sable is known as far south the Kedrovaya river (Kedrovaya Pad' preserve) between
i.e.

Vladivostok and Pos'et (Ognev, 1931),

almost to the state

boundary. The range does not include the steppes of southeastern

Trans-Baikaliya, the steppe areas along the

Onon northward to the lower Ingoda and upper Shilka and eastward approximately to 52
lat.

N.

in the north.

The reconstructed range outlined above

for sable, apparently

was never completely continuous, but apparently, significant gaps were not present in it because of the great ecological flexibility of sable. However, long ago in some places in the 16th century and

"According to Sabeneev (1875, map), the range included a considerable part of bank of the upper Irtysh and across the Alakol'sk lakes (!) it extends to IssykKul'. These rough misrepresentations were based on the extremely confusing and completely false view of N.A. Severtsov (1873) about the fact that in Tien Shan, three marten species existed the pine, stone martens and the sable ("Kashgarsk sable"); however, in this region, they were not yet fully differentiated. The echos of these
the left

views persisted

in literature

almost to the present time.

783
very clearly in the 17th century the sable entirely disappeared in considerable areas both at the periphery of the range (west Siberian forest-steppe and other places) and also within it.

By

the 20th century, this process attained

its

culmination and

the sable range displayed a fragmentation into few more or less considerable sections and a large number of very small ones. All

of them were isolated from each other. At the present time, despite the very great restoration of the
range, as
526

compared

to the

remains fragmented into separate

relatively small.
(Fig. 196).

beginning of the current century, it still parts, some massive and some
are as follows

The regions inhabited by sable

Boundrles of
"Centers" of ^ ccurrence sable
of

524

Fig. 196. General

scheme of recent range of the sable

1955, simplified).

in the

USSR. Figures

designate

the separate "centers"of occurrence. Data up to 1950 (after

Nadeev and Timofeev,

To
and

the largest regions belong: Ural-Ob', occupying the Urals

left bank of the Ob' (Fig. 198); Yenisei, lying along right bank of the Yenisei northward approximately from 57 N. lat.; Sayan,

occupying Sayan and Tuva; Lena between the Lena and Vilyui; Vitim-Trans-Baikaliya lying along the upper Lena, Vitim and along the eastern shore of Baikal; Aldan-Zeisk, situated between the Aldan and lower Amur; Kamchatka, occupying the entire peninsula. There
are

many

smaller regions of less practical importance: Sakhalin,

784

u !

^
U
,S

R " S
00

CJ
.ti

(U

1
?
W)CV
*

* U

if

>>2
J3

<L)

V3

J5

"

^
ii

^2

785

526

Fig.

198.

Boundary of sable distribution in the extreme west of the range Ural-Ob' region of occurrence (after Nadeev and Timofeev, 1955).

in the

Ussuri, southern Trans-Baikaliya and Altai.

are interspersed,
still

Between these centers

smaller and sometimes very small regions of

the au-

sable occupation, sometimes lying very far from each other, as in

Nadeev and Timofeev, 1955; the names of separate centers do not correspond to those given by these
Yakutiya
thors).
(after

Geographic Range outside the Soviet Union

The

(re-established) range includes the northern part of the

Mon-

golian Republic, northeast China, the Korean Peninsula and Ja-

pan.* In the Mongolian Republic, the re-established range occupies Mongolian Altai (sources of Chernyi Irtysh, region southwest of

Kobdo)

in the southeast to the

Bulgun

river (about 92 E. long.),

the region west and east from [Lake] Kosogol, Khangaisk

montane

country south to the upper Orkhon, and westwards including Selenga

and Egin-Gol regions. Apparently, an isolated area of occurrence is found in Kentei south to Bogdo-Ula (a little south of UlanBator), and eastward including the upper Onon and Kerulen. The existence of sable was also recorded in the Mongolian part of the
*Hokkaido only

Sci. Ed.

786
Bol'shoi Khingan [range]. At the present time, sable exists only in Mongolian Altai, Pri-Kosogol'e and Kentei, except for the Onon

and Kerulen.

It is

absent in the Khingans (Bannikov, 1954).

A
lian

finger of the range which extends south along the

Altai,

MongoChinese

apparently

transgresses

somewhat

into

territory.

526

Fig. 199. The course of the natural increase in sable range in the center of occurrence associated within Kuznetsk Altai (upper part of Ob' basin) (after Nadeev and Timofeev,

1955, somewhat simplified).

In northeast China, the natural range of sable occupies almost

all

the country except the

It

Central Manchuria.

exists in the Bol'shoi

Barga steppe and the plains of the former Khingan, Il'khuri-Alin

and Malyi Khingan which extend into the northern part of the country along the Amur, in the mountainous country east of the former Manchuria between the central plains and the Ussuri and

787
528

mountains bordering the Korean Peninsula (Chanbaishan' mountains). At the present time the sable population is strongly depleted, and exists chiefly along the Bol'shoi Khingan and in the northeastern part of the country (Baikov, 1915; Sowerby, 1923;
in the

Lukashkin and Zhernakov, 1934 and others). Details of sable distribution on the Korean Peninsula are not known. In Japan, its
geographical distribution area includes Hokkaido Island, Honshu

(Hondo), Shikoku, Kyushu and Tsushima* (V.H.).

Geographic Variation
Geographic variation
ation
is

in sable is well

marked, and
is

if

taken in

the sense of morphologically extreme forms, the amplitude of vari-

quite significant. This variation

skull

expressed

in general

body and

dimensions

(its

proportions are nearly invariable)

its

and, particularly, colors. Fur quality varies considerably:

density, fluffiness, softness and fineness.

Geographic variations

in sable

practically in the fur trade for a long time,

have been noticed and employed where they were

^^
527
Fig. 200.

Distribution

in

1935

Distribution in

1950

Places of sable introductions

Growth of

sable range in the Ob' -Yenisei section of the range (in part with introductions) (after Nadeev and Timofeev, 1955, simplified).

Populations south of Hokkaido are now usually considered a distinct species, Martes melampus Sci. Ed.

788

Distribution in

1935

Distribution in

1950

Places of sable introductions

527

Fig. 201. Dispersion of sable

and increase in its range in Altai (after Nadeev and Timofeev, 1955, somewhat simplified).

529 eloborated into a detailed

main features orderly and natural system of sable fur "grades". It found its expression in difference in prices of sables from different places, differences that were already established a very long time ago^'^. Serious study of geographic variation in sable from the zoological point of view began only in the 20' s of this century
and
in
its

(Ognev, 1925, 1931), although individual reports appeared earlier (Birulya, 1916, 1918) and attempts at analysis were made more
than 100 years ago (Brandt, 1855).

scheme of variation in the species over its entire range in the USSR, based on significant material was carried out only in the last decade (Kuznetsov, 1941; Nadeev and Timofeev, 1955).
latest
^At

The

the International fur auction in Leningrad in 1961, the highest lot price of

sables of various grades

was

Amur
lars,

31

dollars; Yenisei

Yakutsk

300
1

as follows:

41 dollars, Tuvinsk

dollars,
:

Kuznetsov). Ratio,

14.4. This

and the example

Altai 30 28 Kamchatka 190 dol 67 Barguzinsk 405 dollars per skin hide (B.A.

Tobol'sk

dollars,

dollars,

dollars,

is

a utilitarian demonstration of the reality

of geographic race.

789

528

Fig. 202. Reconstructed species range* of sable

line designates region in Lithuania

Martes (Martes) zibellina L. Dashed and Byelorussia relative to which literature data exist on occurrence of sable in the past. V.G. Heptner.

In the sable very

teristic

wide individual variation

in color is characits

(see above), and therefore the study of

is

geographic

variation

a very difficult task, and racial characteristics are some-

times ill-defined. At the same time, analysis of a mass of collected

which has become possible especially in recent years in connection with growth in the number of sables, showed that individual populations of the species differ from each other in
material,

percentage of various color types.

*Japanese populations south of Hokkaido are

now considered

a separate species,

M. melampus

Sci.

Ed.


790
it was frequently suggested, mainly on the from hunters and traders, that sable color in one narrow region changes depending on the land inhabited (nut mountains) and other pine broad-leaved forest), altitude (valleys of food. Zoologiannual character and even ecological conditions, cally these observations have not been confirmed; however, to completely deny a connection between characteristics and known

In the literature,

basis of information

ecological conditions

is

not suitable. This

is

clear, for

example,

changes in the character of fur in different climates. However, their influence is only revealed over broad areas, for instance in montane and plains countries (see below), but not in individual restricted places as was sometimes indicated (at

from

those

various altitudes, for example).

The present scheme of geographic

variation in sable

was made

on the basis of very extensive material of skulls and, chiefly, large quantities of commercial skins (many thousands). It quite clearly reflects the views of furriers on the types of sable, but, from a zoological viewpoint, still cannot be considered exact. Some
forms are very subtle or doubtful. Though acceptable in the fur business, they do not qualify as representatives of geographic races
in
this concept.

zoology, especially in a stricter and broader interpretation of Some forms were only defined on the basis of neg-

ligible differences in the percentage presence of the three

main

color types, which, as mentioned above, are generally far from

always

stable.

Some forms
it is

will,

undoubtedly, be combined.
an animal dis-

On

the other hand,

necessary to have in mind some special

is

conditions of existence in the sable. This animal

tinguished by

its

great attachment to place within a quite strictly

Its

delineated locality.

range under natural conditions, was always

less isolated parts, especially in the
its

broken into separate more or

east. In the past century, in connection with

extermination,

it

was divided into small parts, surviving

in places for a

long time
influenced
all,

without connection with each other. All this

530 sense,

may have

the formation of small local populations. Easiest of

in this

change

in the ratio

of color types can be expected. In some

for local population

cases, separate

names were noted, apparently,

("colonial") deviations.
Parallel to this, undoubtedly, sable gives entirely defined geo-

graphical forms, irreproachable from a zoological point of view,

and well characterized both by color and meristic characters. Such

791
are the Tobol'sk, Altai, Barguzin, Sakhalin,

Kamchatka and some

geographic forms of sable are found expressed in such a unique index as the value of skins. The skin price of the most expensive category (Barguzin) constitutes
others (see below).

The

reality of

about

1450% of the price of the cheapest (Tobol'sk, see page 788). Some general tendencies may be noted in changes of separate

characters. Thus, the largest sables are encountered to the west

(Ob' lowlands, Altai) and easternmost parts of the range (Kamchatka, Anadyr Territory). Central and southeastern parts are inhabited by sables of moderate and small size. The silkiest fur occurs in the sables from Trans-Baikalia and Yakutia, and the degree of fineness of the fur falls towards the western and eastern
margins of the range and
at the periphery,

manifestation of general regularity is coarsening of the fur in more humid regions, especially in districts

becomes coarse. This mentioned by furriers the

it

of the range and the lightest

plains (Kuznetsov, 1941).

with a maritime climate. The darkest sables inhabit the central part its margins (Ob' lowland, Sakhalin,

Shantar islands). In montane regions, sable are darker than on the

from what has been said above, the scheme of subspecies presented below is preliminary. It is given as material for further use, after Nadeev and Timofeev (1955) and several other sources. The material studied by the first two authors included 856 skulls, 1000 body measurements and 10,100 skins

As

is

clear

examined. Since the dimensions of the different described races of sable are quite similar, dimensions are given of the three extreme forms in the morphological and geographical sense (for information on
the remainder, see Kuznetsov, 1941;

Nadeev and Timofeev, 1955).

Data on weights of sable are given at the end of the list of races. 1. Tobol'sk sable, M. (M.) z. zibellina Linnaeus, 1758. Dimensions large. Fur color light, ranging from gray-brownish
to

tawny-straw.

415-520 mm, females, 370-488 mm; tail length of males, 125-190 mm, females, 115-170 mm; length of hind foot of males, 81-110 mm, females, 75-96 mm; height of ear of males, 45-57 mm, females, 43-55 mm. Weight of males, 1000-1780 g, females, 760-1115 g (Pavlinin, 1963; from exten-

Body

length of males,

sive original data and literature).

85.7-90.6

Condylobasal length of skull of males (159 samples) 79.7-M mm, females (87) 73.8-M 78.9-83.8 mm; zygomatic

792
width of males, 44.1-M 48.0-54.8

mm;

height of male skull

mm, females 39.0-M 43.6-47.2 29.5-M 32.4-35.0 mm; females 27.4-

M29.8-32.2 mm (Pavlinin, 1963). Golovka and Podgolovka sorts are absent, Vorotovaya constitutes 40% and Mekhovaya 60% [see pp. 763-764]. Found in Pechora basin. North and Middle Urals, western Siberia to Ob' river. Absent outside USSR.
Lightest colored of
all

our sables.

It

is

suggested that the

Pechora or Pechora-Ural sable differs from the Trans-Ural. At the same time, it is possible that this form occupies West Siberia as far
as the Yenisei.
2.

Kuznetsk sable, M. (M.)

z.

tomensis Nadejev

et

Timofejev,

1955.

Smaller than Tobol'sk sable. Color considerably darker than Tobol'sk form and somewhat darker than Angarsk, but lighter than
Altai.

Golovka and Podgolovka sorts constitute 19%, Vorotovaya type 41.9% and Mekhovaya 46.1%. Found in the western slopes of Kuznetsk Altai, Tom' [river]
system.

Absent outside USSR. 3. Altai sable, M. (M.)

altaica).
531

z-

averini Bashanov, 1943

(synonym

dimensions comparatively large, color quite dark and varies from sandy-yellow to dark brown. Golovka and Podgolovka sorts 26%, Vorotovaya 66% and
skull

Body and

Mekhovaya 8% (n=1000). Found in Altai, particularly Bukhtarma of Tuvinsk ASSR (Kemchug basin).
Outside

basin, northeastern part

USSR, found

in

southern slopes of Mongolian Altai

(Bulugun

river).

In several places in Altai,

Barguzin sable was introduced, yieldjeniseensis Ognev, 1925.

ing hybrids with local sable.

4.

Yenisei sable, M. (M.)

z.

Similar to Tobol'sk, but differs in smaller skull dimensions

and darker fur color. Fur

is

loose, but very coarse.

Found

in interfluve

of Ob' and Yenisei rivers from northern

border of taiga southwards to

Chulym

river.

Absent outside USSR.

793
5.

Angara

sable,

M. (M.)

z.

angarensis Nadejev

et

Timofejev,

1955.
Differs from Yenisei in smaller average skull dimensions.

General color comparatively

Tobol'sk and Yenisei.

light,

but noticeably darker than

Golovka and Podgolovka sorts constitute 9.7%, Vorotovaya 77.8% and Mekhovaya 12.5% (n = 2600). Found from Yenisei valley eastward to Oka (tributary of Angara river), along right bank of Angara northward to watershed with Podkamennaya Tunguska and southward to Kansk steppe. Absent outside USSR. 6. Sayan sable, M. (M.) z. sajanensis Ognev, 1925. Differs from Angara by considerably darker color. General tone of fur tawny-brownish with admixture of pale-straw underfur. Golovka and Podgolovka categories constitute 17.4%. Vorotovaya 70.9% and Mekhovaya 11.7% (n = 2679).

Found
7.

in

Sayans.

in Kosogol lake district in Mongolia. Tungusska sable, M. (M.) z. tungussensis Kusnetzov, 1941. Differs from Tobol'sk in less massive skull, somewhat shorter braincase and considerably darker fur. Golovka and Podgolovka sorts constitute 6%, Vorotovaya 55% and Mekhovaya 40%. Found in basins of Podkamennaya and left tributaries of Nizhnaya Tungusska. Absent outside USSR. 8. Ilimpiisk sable, M. (M.) z. ilimpiensis Nadejev et Timofejev,

Outside

USSR, found

1955.
Similar to preceding form in body and skull dimensions but
differs

by its considerably darker color. Golovka and Podgolovka sorts constitute 16%, Vorotovaya 53%,
to northern border of

and Mekhovaya

31% (n = 2600). Found north of Nizhnaya Tungusska

species range, and eastward from Vivi to Lena, southern border

between 64 and 64 N. lat. Taiga of Tura, Kureika, Kotyi, and Anabar basins and upper reaches of Olenek. 9. Barguzin sable, M. (M.) [z.] princeps Birula, 1922 (syno-

nym

baicalensis)
relatively small

Dimensions

one of smallest races of species;

lighter than

color entirely dark, but overall

somewhat

Vitim sable.

794

Body

length of males,

382-410 mm; tail length 120-M 129-145 mm; condylobasal length of males, 73.9-M 77.981.0 mm, females, 68.4-M 73.1-79.1 mm; zygomatic width of males, 39.6-M 41.6-44.5 mm, females, 37.7 M-39.7-41.8 mm; height of male skull, 29.6-M 30.5-32.3 mm, females, 28.5-M 29.731.4
532

394-M 415-420 mm, females, 360-M of males, 122-M 144-155 mm, females,

mm.
sorts constitute

Golovka and Podgolovka

and Mekhovaya

45%, Vorotovaya 46%

9%

(n=786).

Found on eastern shore of [Lake] Baikal; Barguzin range from

Barguzin to watershed with upper Angara. Absent outside USSR. One of the most valuable forms. 10. Vitim sable, M. (M.) z. vitimensis Nadejev et Timofejev, 1955. Larger than Barguzin, but smaller than Tobol'sk, Altai, Kamchatka and Chikoisk. Color is darker than in Barguzin, very dark, intense blackish-brown. Throat spot weakly defined. Fur
right tributaries of
fluffier

and silkier than Barguzin. Golovka and Podgolovka sorts constitute 56%, Vorotovaya

42%

and Mekhovaya 2% (n=5529). Found on right banks of Kirenga and Lena, upper Vitim and upper Angara and northern extremity of Baikal. Absent outside USSR. Form closest to Barguzin. One of most valuable. 11. Chikoisk sable, M. (M.) z. obscura Nadejev et Timofejev,
1955.

Darkest form, considerably darker than Barguzin.

One

of large

forms, inferior only to Tobol'sk, Altai and Kamchatka.

Golovka and Podgolovka sorts constitute 71%, Vorotovaya 27.6% and Mekhovaya 0.9% (n = 483). Found in southeastern extremity of Yablonovoi range Chikoi

system, the right tributaries of Selenga and Ingoda.

Outside USSR Kentei [mountains] Extremely valuable form.

in

Mongolian Republic.

12. Yakutsk sable, M. {M.) z. jacutensis Novikov, 1956. Dimensions particularly small, smallest race of species in USSR.

Pelage very fluffy and silky. Color strongly variable, but the dark
sables prevail.

Golovka and Podgolovka sorts constitute 53%, Vorotovaya and Mekhovaya 5% (Kuznetsov, 1941).

42%

795

Found

in

upper Aldan and Zeya eastward

to

Uda.

Absent outside USSR. For naming of form, see synonymy of species. 13. Sakhalin sable, M. (M.) z. sahalinensis Ognev, 1925. Dimensions very small, color very light with predominance of
reddish
tints.

Golovka and Podgolovka

and Mekhovaya

sorts constitute

4%, Vorotovaya 59%

47%

(Kuznetsov, 1941).

Found
It is

in

Sakhalin [Island].

Absent outside USSR.

probable, the

according to some data,

name brachyura belongs to the form since, Temminck described this form from skins

obtained from Sakhalin. Relationship of Sakhalin sable to that of Hokkaido {brachyura) is not clear. 14. Ussuri sable, M. (M.) z- arsenjevi Kusnetzov, 1941. Dimensions as in Sakhalin sable, color medium dark, pelage
less fluffy, as if clipped, but soft

and

fine.

Golovka and Podgolovka and Mekhovaya 17%.

sorts constitute

16%, Vorotovaya

67%

Found

in

Ussuri Territory, mainly in Sikhote-Alin' taiga.

in

Outside
in

USSR, found

contiguous parts of North-east China.

Relationship of this form to form coreenis {hamgyensis) found

Korean Peninsula needs clarification. 15. Shan tar sable, M. (/.) z. schantaricus Kusnetzov, 1941. Dimensions as in Ussuri sable. Color darker than Sakhalin and
sorts constitute

Ussuri. Pelage quite fluffy.

533

Golovka and Podgolovka and Mekhovaya 5%.

32%, Vorotovaya 63%

Found in Shantar Islands, the basin of lower course of Amur and Tuguro-Udsk coast of Okhotsk Sea.
Absent outside the USSR. 16. Kamchatka sable, M. (M.) z. kamtschadalica Birula, 1918 (synonym kamtschatica). Distinguished by large dimensions of body and skull largest form of species. Color strongly variable, but dark individuals pre-

dominate. Pelage very thick, but insufficiently

soft.

Body length of males, 445-M 490-580 mm, females, 415, M 448-510 mm; tail length of males, 147, M 160-175 mm,
females, 132,
85.6,

152-172

90.4-94.6

mm

mm; condylobasal length of male (M 88.5 mm-93), females, 79-4, M

skull,

82.1-

796
84.4

mm

(
mm

80.8

mm-76); zygomatic width of males,

50.2 mm-81), females, 43.6,

51.3,

53.5-56.2

(M

46.3-52.8

mm

mm-74); skull height of males, 31.6, M 33.5-35.1 mm (M mm-94), females, 29.8, M 31.5-39.6 mm (M 30.6 mm-75) (Timofeev and Nadeev, 1955, from 56 skulls; in parentheses, mean and number of individuals from Kuznetsov, 1941).

(M

44.9

33.3

Golovka and Podgolovka

sorts constitute

16.0%, Vorotovaya

76.0% and Mekhovaya 8.0%. Found in Kamchatka and Koryak land

Absent outside USSR. 17. Kuril sable, M. (M.)
sufficiently clear.
z.

in the

Apuka

basin.

brachyura Temminck, 1840.

in the Kuril Islands are
its

Diagnostic features of sable found

not

Some
all

authors relate
it

name

to the

named

Japa-

nese form. Apparently,

belongs to the Sakhalin

in sable type.

On

the whole,

forms enumerated

may be

regarded as aggre-

gating into seven groups (large populations): Tobol'sk

Urals and West Siberia to Yenisei; Altai [and]; Sayan

ASSR, Sayans and

north to Kuznetsk Altai;

of Yenisei to Lena; Trans-Baikaliya

range, east to watershed of

south

Pechora, Tuvinsk Yenisei bank

right

to

border of species

Zeya and Bureya,

north, including Vitim;

Is-

lower Amur, Sakhalin, Ussuri Territory and Shantar lands; Kamchatka Kamchatka and Koryak land (B.A. Kuznetsov).
Sakhalin

Weight of sables from various

females (125) 880,

parts of the range is as follows

(Kuznetsov, 1941): Yenisei sable males (93) 1030,

1307-1500

Sayan sable July males g, 980-1300 g, 1433-1800 g, females (21) 870, (22) 1200, 1472-1810 January males (22) 1110, g, females (21) 820, 1239-1600 1132-1560 g; Sakhalin sable July males (22) 940, g, females (25) 800, M905-1130 g, January males (22) 880, 916-1120 g; Kamchatka sa1193-1500g, females (25) 740, about 1500-1869 g, females (5) 600, blemales (4) 800, about 1100-1140 g. Outside the limits of the USSR, the following forms are noted (sometimes, several of them are related to "/. melampus"): 1) M. in Hokkaido, Japan; 2) M. (M.) z. brachyura Temminck, 1844 melampus Wagner, 1840 in Honshu (Hondo), Shikoku, z. Kyushu Japan; 3) M. (M.) z. tsuensis Thomas, 1897 in Tsushima Island in the Korean strait; 4) M. (M.) z. koreensis Kuroda at Mori, 1923 (synonym hamgyensis, Korean Peninsula) (V.H.).

1182-1250

g;

M M M

797
Biology
Population.

By

the 30' s of the 20th century the

number of

sable

had become very small as a result of their rapacious destruction for about a whole century; at the present time their numbers are continuously increasing, as a result of the system of state measures for
534 protection

and they

far

and restoration of this very valuable fur-bearing animal exceed the prerevolutionary.
in Siberia where which previously

At the present time, there are many regions

the population density of sable attained a level

seemed impossible and was not actually

were there
territories

present.

The

latter is ex-

plained in that nowhere in the past within the range of the sable,

which were released due

to scarcity of sable

hunters. In connection with this, in

many

places, areas

were over-

populated by sables that led to undesirable consequences. Such

saturation could only occur in the period before the appearance of

the fur trade.

The present number of sable has already for some time allowed the establishment of a planned, licenced harvest. Sable are
quite unevenly distributed within the limits of their range. This

condition

is

due not only

to different degrees of favorable environ-

mental conditions but also to the fact that in the different parts of
the range the active measures for restoration of
its

begun

at different times.
still

Therefore,

at

present the

numbers were number and

density of sable

do not correspond everywhere

distribution
is

to the capacity

of taiga areas and

lation.

its

not the result of natural regutotal

Evaluated on the results of the hunting harvest, the

in the

number of sable

USSR

could be very provisionally calcu-

lated to be about 500,000.

For the greater part of the present range, the average density
does not exceed
1.5

animals per 10 km^ (Sverdlovsk, Tyumen,

and Chita oblasts, Buryat ASSR, Kamchatka). In places, this density is considerably lower (Omsk, Tomsk, Amur oblasts, Tuvinsk ASSR and others). In individual regions (Eastern Sayan, Altai and others) the average density reaches 2.5-5.0 and 10-12 individuals per 1000 hectares (Nadeev, 1961). In southern

Kemerov,

Irkutsk,

Altai, in areas of highest sable density,

one sable has 3 km^ (3.3

sables in 1000 hectares), while in the areas that are regularly exploited

4,2

km^ (Afanas'ev,

1961).

798
In

Kamchatka,

in the best areas (forests

of Erman's birch

[B.

ermani], mixed broad-leaved-fir-birch forests), density reaches 1.52.0 sables per 10

km^

with 8.3-8.5 track encounters per 10

km

in

good areas
ests)

(forests of white birch [B. alba], flooded forests)

1.4

(6.8-7.3 track encounters), in moderate (broad-leaved and

fir for-

1.3 (4.8 track encounters)

and
is

in the

worse

0.1-0.3 (0.8sables
is

1.4 track encounters).

The

total

number of Kamchatka

estimated

at

19,000 (average catch

7,650 individuals, equaling

an average growth of 40.3%; A. A. Vershinin).

In northwestern Yakutia, into consideration uninhabited regions

one sable has 15-26 km^ and taking 100-120 km^ (Tavrovskii,

1958).

However, contemporary census methods are unreliable and often

extrapolated over large areas with insufficient basis.
Habitat. Sables inhabit large massifs of northern coniferous

They are in equal measure native to montane and plains taiga (urman)*, penetrating to the zone of sparse tree growth in the north and in the mountains. They do not go beyond the limits of coniferous taiga. They avoid cultivated
forests of the taiga type.
areas,

and cross
is

their boundaries only during times of migration.

Preference

shown

for dark coniferous plantations almost always

is

having

in their

composition Siberian stone pine. This

because of

the great availability of food (pine-nuts, forest voles) and the protective nature of this plant formation.

Only

as a result of irrational utilization, sable

talus
in the

inhabitants of

montane taiga, rocky montane balds. At the present time,

where lowland pine
forests

became mainly and frequently even

same way
as in the

distant past, sable willingly occupy and attain considerable density

and swamps predominate, although even

belts of dark coniferous forests

here, they prefer the islands of nut-pine and spruce forests, mar-

gins of
535

moss bogs and narrow

forests of

along river banks. Sable also lives under unique conditions such as

montane

Erman's birch

in

Kamchatka,

in larch forest

with an understory of green alder [. viridis] in the Zhigansk

region and even in forest tundra in the Gizhiga and Penzhina regions.

The close linkage of

sively conditioned

sable with the Siberian nut-pine

is

excluit

by the general scarcity of food available for

*Local word

Sci. Ed.

799
in the Siberian taiga. This

connection

is

not an indispensable con-

dition of

its

existence.
is

Food. The sable

a polyphagus carnivore. In addition to

warm-

blooded animals, plant food seeds of Siberian and Korean nutpine and prostrate nut-pine, berries and insects are of great
significance in
its

in various places.

food. The composition of its food varies greatly For example, differences in food type of lowland
in pine marten, sable foods

and montane sables are quantifiable. As

The larger and stronger males often consume larger prey capercaillie, blackcock, and white hares. Such differences in nutrition were noticed in the foothills of the East Sayans (Dul'keiti, 1956). They are also confirmed in the northern Urals (Zaleker and Poluzadov, 1955) as well as for the Yenisei sable (Kiris, 1953). However,
reveal geographical, stand-type and sexual differences.

several investigators did not find these differences (Raevskii, 1947;

Lavrov, 1953). This can be explained easily

in

that in

many

Fig. 203. Habitat of sable in sparse

growth of montane nut-pines

at the

upper limit

of forest. Altai preserve. Photograph by G.D. Dul'keit.

800
regions, large food objects are absent or are

met with

rarely

and

sporadically.
In the northeastern Pribaikal,
series of other regions, only

nuts

or

the forest

Khabarovsk Territory and a two elements forest voles and pinevoles and cowberry [V. vitis-idaea] are of

essential significance in sable nutrition.

The geographical and seasonal changes

in sable

foods are well

manifested (see Tables 50, 51). For Kamchatka, the willow ptarmigan, root vole, "nuts" of prostrate nut-pine and fish are character536 istic

foods. Red-backed voles, pine "nuts" and mountain ash

in

predominate

foods of the Barguzin sable. For Kondo-Sos'vinsk

for

sable, capercaillie and, in general, tetraonid birds are typically the

most often encountered, and

cowberry.

Pri-Amur sable

forest voles

and

Fig. 204.

Taiga along

Mana

river

a tributary of

tiie

Yenisei, habitat of sable. "Stolby'

preserve south of Krasnoyarsk. Photograph by G.D. Dul'keit.


801

The food composition of sable changes in different years depending upon yields. With failure of the plant food crop, the role of warm-blooded animals significantly increases and vice-versa (Shaposhnikov, 1956). A lessening of the role of one of the foods
causes increased consumption of another. Usually, a deficiency of

one or two main food is compensated by the ary foods, or those which are not regularly
537

utilization of secondutilized.

Food composition
is

is

not the

same even between small

populations living in separate montane valleys in one region. This

well illustrated by data from the Barguzin preserve (N.M.

Baevskaya) (Table 49).

Table 49. Food composition of populations living along different rivers in Barguzin preserve (occurrence in %)

Food composition
Mouselike rodents
Birds
Reptiles
Insects

Bol'shaya river
90.0
13.0 12.2

Davshe
75.0
7.5

river

Tarkulik river 92.3

3.8 3.8

5.0

23.6 30.0
16.9

25.0 55.0 20.0

15.3

Pine "nuts"
Berries

73.0
3.8

Food composition of
The seasonal

sable changes in

snowy and nonsnowy

periods of the year in various parts of the range (Tables 50, 51).
variations in sable nutrition are not as sharply

manifested as in the pine marten. They are determined by the

seasonality of a series of foods

insects, berries,

and

nuts,

which

can serve as food for sable only

in the

warm

period of the year. In

Kamchatka, fish becomes available for sable only in the salmon spawning period. On the other hand, seasonality determines the
availability of food for sable; therefore, in the snow-free period,

forest voles, especially gray [C. rufocanus],

schisticolor], shrews

wood lemmings [L. and Siberian chipmunk are significantly more available than in the snowy period. In this sense, winters with deep snow and those with little snow differ from each other. In winters
with deep snow, sables more frequently catch tetraonid birds

capercailles and hazelhens

In

when they spend the night under snow. winter periods with deep snow, sable feed mainly on berries of snow on autumn, or when there mountain and
ash,
in
is little

cowberry, and
ficult to

summer, on blueberry. Deep snow makes obtain voles, pine "nuts" and some other food.
in

it

dif-


802
Table 50. Geographic variation in sable foods in the snow-free period of the year (occurrence in%).

Region,

West
Siberian

Northeastern
Altai

Barguzin
preserve

Middle
Sakhalin

\
\

Number
items,

of

lowlands

\ Investigator

76

Food

\
\

Raevskii,

Shaposhnikov,

composition
Forest voles

1947
57.7

1956
36.8
3.9
1.3

314 Timofeev 1948

73.5
3.5 2.5

E.

357 Volkova

Siberian

chipmunk
squirrel

Eurasian flying-squirrel

Common
*Pika

7.7
7.7
23.1

64.0
5.0
1.0

3.9
0.5

1.9

No No

0.2
2.0

Shrews
Birds

19.7
1.0

13.0
6.9
1.2 1.2

25.0
information

Tetraonids

7.7 7.7 3.8

Hazel hen
Bird eggs
Insects

3.9

information
0.6

1.3

0.6 5.7

53.8
3.8

13.1

15.0

Pine "nuts"
Berries

36.8
13.1
is

37.6
35.7
used, rather than the

34.0

44.5

66.0

*The Russian word senostavka ("haystacker")

more com-

mon pishchuka

Sci. Ed.

this explains, for

Sable eat carrion and the food remains of other carnivores example the occurrence of squirrel in the food of
Utilization of the Eurasian flying-squirrel [Pteromys volans]
is
it

sable (Shaposhnikov, 1956; P.P. Tarasov).

associated with diurnal activity of sable; the latter can overtake

in its holes

only during the day (the flying squirrel

is

a nocturnal

animal).
In

ponent

voles varied

Kamchatka, the frequency of the most important food comin different years from 35% to 84% (A. A.
the factual data on sable foods, one

Vershinin) (Table 52).

Summing up

may
food
its

affirm
is

that the significance of the

Siberian chipmunk

in its

ex-

tremely limited and

is

not found everywhere: in summer,

occur-

rence in sable food does not exceed

less.

3-5% and

in winter, is

even

The old idea about

the great significance of haystackers (pikas)

in sable nutrition is not

confirmed. In winter foods, the share of

539

pikas (by percent of occurrence) ranges from 0.9 to 10.1%, more usually not exceeding 5%. In summer it is still less. Pikas occur in
sable foods in only

46%

of the regions where

its

nutrition has

been

!2

tN

,
I

-^ OS

\ 1^
t- fN <N

-5

*
'-:

ZZ ZZ
rt
1 1

-:

IS

2
I

OS

122

rt vo

S -

P M
OS

3
<5
'

S
00

;:::

^
00
rsl

(M

00

<

OS OS

OS
1

->

>

00
I

-^

-i

Z
J2

>-

3
W 00 1 ^ UI ^ ! ^ 3 -^ Q
'^

r^ cs

1?^

t^

(
on

1^

q
>
g (

r-^

1^

r^ so
I I

OS so

OS

^ ooro

^
^

'*-'

osmiTi

vo

Tj-

inO>n-^rslOO(N

OS (N

w-l

> S ^ ^ 2i 3 Q
n^
'*3

~
00 <N
I

t-;

* -^

SO OO

00

:2
OS t~

(N

OS OS fN so iri in

1/-1

04

csi
I

n OO
I

(N
I I

-; --^ fS <N N

00 Tt

r<-)

I I

1 00

^^

-Z ZZ

=
._
t"

HHH

>> fS -^ SO

*->.'*
4)

2
{^

fc

(U

.t;

^
CQ

5^ 4

;
N

(JuOooooOion^m

Ji

J-

1)

" % 3

..

804
Table 52. Results of a multiyear study of materials concerning sable nutrition in Kamchatka (1409 stomachs; A.A. Vershinin)

Food composition

Occurrence

(in

%)

Specific weight of
individual foods

Mammals
Voles

69.8
61.1

66.6

61.0
24.0
12.7

Root voles [Microtus oeconomus] Red-backed voles {Clethrionomys Red-gray vole [C. rufocanus]
Squirrel [Sciurus vulgaris]

24.0
rutilus]

20.4
16.5

19.4

0.8
2.7

0.4

White hare [Lepus timidus] Pika [Ochotona hyperborea]

Birds
Capercaillie [Tetrao parvirostris]

0.4

2.4
6.1 1.2 1.2

20.8
1.7
1.5

Willow and rock ptarmigan

Small birds
Bird eggs

12.5
1.1

Fish

18.1

Salmon
Insects

10.2
2.5
insects

Hymenopterous Plant food

Berries

0.9

11.9
15.2
2.7

3.6

53.5

"Nutlets" of prostrate nut pine

10.3

12.5

Mountain ash [Sorbus] Hawthorn [Crataegus] Chokecherry [Prunus sp.] Rose [Rosa sp.]

21.6 20.9
4.8
7.3

3.6

6.1

Studied.

The percentage of
it

squirrel occurring in winter foods of

sable ranges from 0.2 to 11.3%, averaging 3.5%. In the

period,

summer

ranges from 0.2 to 7.7% (on average 2.9%).

As
rel in

a result of data

sable nutrition (Timofeev, 1962)

communicated on the significance of squirit was revealed that in various

in various years, the

parts of the range

and

percentage of occurpercentage of

rence of squirrel in stomachs and extrement of sable varied from

and 26%; on average, 4.4%. Of 33 data

squirrel occurrence

series, the

was more than 10% in only 6 cases. It may be considered that: 1) The role of squirrel in the diet of sable in different years and seasons is not the same and is not connected with degree of squirrel abundance in areas inhabited by sable; 2) this role increases in the periods of scarcity of other main foods of sable (voles, pine "nuts"); 3) sables catch squirrel mainly on the ground; pursuit of squirrels in the tops of trees end unsuccessfully in most cases; 4) destruction of small squirrels in the nest is not


805
yet proven, 5) the influence of sable on reduction in
squirrels
is

number of

not so great that

it

can explain reductions in the result-

ing squirrel harvest. These far from complete data indicate that
squirrel
is

of secondary importance or even an accidental element

in sable nutrition.

The
inside

sable cannot penetrate into the burrows of pika and chipto its large size. In winter, the

munk due
its

chipmunk

sleeps deep
it

burrow, and even the brown bear cannot dig

in

out.

However, same way as burrows with

summer
it

sables catch

chipmunks by ambush

in the

does with pika. In winter, pikas come out of their

540

hay stores under the snow where they are sometimes overtaken by sable. Sable catch tetraonid birds in their snow burrows, but this does not occur often. In the north Urals, the sable and kida catch one tetraonid for every 10-15 attempts, when the
their

birds succeed in flying away.

In captivity,
it

all,

but hid

when offered 200 to 300 g food, sable did not eat some (Dul'keit, 1929); a norm of 170-180 g and

150-160 g of food were sufficient for males and females respectively. Under natural conditions, pine marten and sable do not eat more than 1/10 of their live weight per day, usually 100-170 g (P.B. Yurgenson; Timofeev, 1951). They usually leave food behind, when food can be obtained in abundance. Concerning the statement (Petryaev, 1950), that sable eat in one day an amount of food equal to half its live weight, we consider it impossible. The volume of the sable stomach does not exceed 120 to 200 cm^ (Sokolov, 1941), and in nature a full stomach was practically never found in sable. In Kamchatka, the maximum weight of stomach
contents in sable was established as 36 g with an average of 24.4 g. In that study, in one stomach up to 6 voles, 500 "nutlets" of

dwarf pine, 160 fruits of mountain ash, 50 rose, and up to 300 hawthorn, were found at one time. Differences in the sable diet may be observed in individuals of different sex: in females, the percentage of empty stomachs is 56.6 and in males 52.7, and the average weight of stomach contents is

22 g and 26.4 g respectively, and fatness

in

females

is

also less.

Young

(yearling) sables also feed less well. Sable diets differ in

separate regions of
Vershinin).

Kamchatka and

in

different years

(A. A.

prey,

The sable does not make food caches. it makes a temporary lair nearby.

In the presence of large

806

Home

range. In August, the sable

litter

disperses.

Young
range.

sables begin to wander, searching for a permanent

home

Food shortage
settlement
is

intensifies their wandering.

When

the period of

completed, sable become

is

strictly

sedentary animals.

The home
until death.

range selected

usually used for

many

years, often

Only

forest fire, felling of the forest

its

and similar events

make

the sable leave

home

range. After the death of the "owner",

its boundaries which by the boundaries of the home ranges of neighboring sables (Raevskii, 1947). Within each range, only one animal, or a female and her litter, live. With significant population density, however, neighboring home ranges often impinge on one another. Occasions have been observed when the owner of a home range pursued and drove away from its boundary another sable that had

another one occupies the range, preserving

are determined

penetrated thence. Rarely, fights

may

also occur, usually during

low

densities.

More

often, sables avoid transgressing into the range

of their neighbor, limiting themselves to their peripheries.

In case of high density,
it

seems
is

as

though the reflex of range

popu-

defense

is

lost.

Competition

not intensified, but migration

less settled part of the

appears, mainly
lation (O.K.

among

juveniles

the
home

Gusev").

The

size of the

home range

is

determined by the abundance of

range area of various quali-

food, the presence of escape and nesting conditions, and also the

percentage correlation inside the

tative factors including those

which are not all necessary for the sable. Each home range seems to be divided into separate sections of daily activity which are periodically visited by the sable. Sometimes, within the boundaries of such a range
it

will stay for

some

days, whereas others are not visited for a long time.

The

areal

dimensions of the home range, equally with the length

of the daily movements, are the best criteria (bonite) for the
quality of sable range. In the

American sable [= marten, Martes

541

americana] length of daily movement in the USA is equal to 8.4 km on the average, varying from 3.2 to 14.4 km. Correspondingly, the range of daily activity equals 248 ha on the average, varying from 129 to 518 ha (Marshall, 1951). By periodically changing
^^There exists the opinion (Raevskii, 1947) that excess individuals inhabiting the
central parts of a densely settled territory die as a result of intensive competition since

such sable do not have the rank to take over ranges inhabited by other individuals. Later (Tavrovskii, 1958), it was shown that this opinion is mistaken.

807

541

Fig. 205. Littered

montane

taiga in the "Stolby" preserve (Western

lair

Sayans south of

Krasnoyarsk). Place of day-time

of sable in areas where Barguzin sable was

introduced. Photograph by G.D. Dul'keit.

these ranges, one animal

may occupy

home range

of 2590-3885

ha

(states of

Idaho and Montana). The size of the daily range of

in

233 ha on average for males (77.7-440.3 ha) and for females 69.9 ha (7.7181.3 ha). Through repeated tracing of the tracks of one male, daily length [of the track] was about 0.8 km, varying from 0[.0] to 12.5 km. As in Siberian sable, the borders of the range usually intersect each other (Vernon, Hawley, 1957)*. In Kamchatka, the size of the home range rarely exceeds 10 km^ (Plechev, 1939), on Bol'shoi Shantar Island, on average 13 km^from 5 to 30 km) (Dul'keit, 1929). In the Zhigansk region

American marten

Glacier National Park (Montana)

is

(Yakutiya),

home range comprises about 30 km^ (Romanov,

Hawley, V.[ernon], 1957

1938).

In northwestern Yakutiya (Tavrovskii, 1958), 5 out of 8 sable trails

*Correct citation
is

Sci. Ed.

808
did not exceed a circle 4
eastern

km

in

diameter (about 12.6 km^),

in

km^ (Kozhanchikovy, 1924), on the about 10 km^ (Doppel'mair, 1926), northeastern shore of Baikal (Altai) from 8 to 12-15 km^ (V.V. river in the upper Katun'
Sayan
to 10

from 4

Dmitriev and P. Bashilov), in the former Kondo-Sosyvinsk preserve, on average about 10 km^ (Raevskii, 1947), and in the north Urals on the upper Ilych river about 4-6 km^. In the Eastern

moderate crop of principal food, a home range of 2.5-3.0 km^ is sufficient for one sable (Dul'keit,

Sayan

foothills, in case of a

1957); the

maximum

density in this case was

sable per 2.6-2.7

kml
Within the limits of the distribution
quently, there
is

that are observed, conse-

fluctuation in size of range

from 4

to

30 km^.

542

Such fluctuations occur locally and in limited territories. Burrows and shelters. Breeding, winter, autumn and temporary dens may be distinguished (Raevskii, 1947). Breeding and winter dens are permanent. They are usually in the form of dry
hollows in trees and stumps and in logs that are not completely in holes in the ground and lying on the ground, and quite rarely in crevices in rock slides. The nesting chamber is covered with dry

grass, lichens, wool, feathers,

down,

etc.

Not much bedding occurs.

/
Fig.

206. Barguzin sable in a

snow

lair.

"Stolby" preserve south of Krasnoyarsk.

Photograph by G.D. Dul'keit.

809

Winter dens are permanent only where winter

is

distinguished

by

its

severity. In

Kamchatka (V.A. Kotov),

in northeastern Altai

they are not manifested. Permanent winter nests

significant

(Shaposhnikov, 1956) and in Primor'sk Territory (Bromlei, 1956), show signs of

occupation
others).

(ice-covered

entrances,

excrement,

ectoparasites and

Autumn

lairs are

characterized by less

evidence of occupation: a tree hole. For permanent habitation in winter they are very wet or cold. Temporary and occasional lairs

boundary of the home range, the Sayan foothills, 51% of sable dens in winter were under windfalls, roots and uprooted trees, and 49% in cliffs, rock slides, and boulders (Dul'keit, 1957).
are of various forms. Within the

sable has

some of

these. In the Eastern

Daily activity and behavior. The sable belongs among those animals with an indefinitely defined rhythm of daily activity. Individual sables are observed with crepuscular-nocturnal
diurnal types of activity.
It is

others with

not even rare that one and the same

is active during the day, and then in the twilight and at depending upon the success of its hunting, weather conditions and other events. In the northern taiga, in connection with the characteristics of the light regime in the course of the year, the daily rhythm of sable life and its victims is particularly unclear (Raevskii, 1947). In a series of cases, change in the predominant

animal

night,

types of daily activity have taken place as a result of intensive

pursuit

by hunters, who can more

easily extirpate animals with a

crepuscular-nocturnal type of activity. In specific cases such changes

in

type of daily activity of a population over a 100-year period

may

be established by comparing recent data with these of L.P.

Sabaneeva (1875). There is no single rhythm of daily activity in different populations. In the northern and middle Urals and on the shore of
543

Baikal, the diurnal prevails, and in the Trans-Urals

noctural.

crepuscular-

On

Blol'shoi] Shantar Island and in the northeastern Altai,

the animals have

no definite rhythm. In the

foothills of the Eastern

Sayan, in
1957).

68%

of the cases, sables hunt during the day (Dul'keit,

The sable
not capable of

is

a typical terrestrial carnivore.

it

On

trees,

it

does not
is

feel self-confident,

climbs more poorly than pine marten and

directly

from one tree to another as can the latter. Having climbed a tree, it jumps down again. The sable is well adapted to living on the ground surface in severe snowy

moving

810

543

Fig. 207. Sable in natural habitat. Stolby preserve south of Krasnoyarsk.

Photograph

by G.D. Dul'keit.

winters.

It

orients itself well within the subnival space,

moving
is

about freely there and capturing food items. The senses of hearing and smell are well developed. Vision

weak and monochromic. The

In
its

sable

swims

well.

search for food during a 24-hour period, the sable jour-

neys a quite considerable distance. The length of the daily winter trail depends on the abundance of food and the degree of its availability to the sable, as well as

on the type of snow cover and sex

of the animal (Dul'keit, 1957). While moving on soft snow, the

sable quickly
that with the
tires. The small size of the female leads to the fact same number of jumps and energy utilized, the fe-

male runs
sable

a shorter distance than the male.

In the basins of

Konda and Sos'va

rivers (Raevskii, 1947), the

moves 2-3

km

per day (early in spring, 4-5 km); in the

northern Urals (Ivdel'skii region)

(Shaposhnikov, 1956)

from

8-9
km

km;

northeastern Altai

3 to 11.5

km

(average, 5.7). In the

southern Altai (Kazakhstan), the length of daily trail of the sable ranges from 0.2 to 5 km, usually 3-4 km. During a snowfall, the
sable rarely

moves more than 2

(Afanas'ev, 1961). For the

is

Eastern Sayan (Solov'ev 1921), the path

5

indicated to be about

km; in Barguzin preserve about 10 km on the average; in Kamchatka (Kazarinov, 1946) about 12-15 km; in the Sikhotefrom Alin' range (Bromlei, 1956) and in snow 12-14 cm deep

811
1.3 to 3.1 km, averaging 2.4 km. In the foothills of the E[astern] Sayan (Dul'keit, 1957), length of daily sable trails in the 19531955 period did not exceed 9.2 km. Average length of the daily 4.5 km. Average trail was 5.6 km for males, and for females length of the daily trails (both sexes) was 5.1 km. In the first half

of winter,

when

it

is

easier to capture

its

food,

it

runs 4.3

day, and in the second half of winter

sable, the longer
544 as well as
its trail

km. The larger the per day. Deep snow and frost shorten it,
in years of nut pine harvest, the sable

5.9

km

per

abundant food;

sometimes becomes satiated in a 1-km journey. In northwestern Yakutiya (Tavrovskii, 1958), the average length of the daily journey of the sable is about 4 km. A circular route is characteristic for sable; along its way it rarely doubles back or turns aside. Even while fattening, the sable's path is distinguished by its characteristic

directional integrity. In winter, the sable catches mouse-like

rodents under the snow; on

warmer days, they often ambush them

Timofeev,
It

on the surface of the

snow

like a cat (northeastern Altai:

1951; Shaposhnikov, 1956).

catches tetraonid birds in the

It

snow

holes where they spend the night.

searches for nut pine "nutlets"

and cowberries under the snow. Mountain ash berries are eaten
directly

from the branches.

is

Seasonal migration and transgressions. The sable

tary animal, but translocations

a seden-

do occur in the species. Above all, dispersal of young in August to a new place, there is the yearly beyond the boundary of the home range where the litter was weaned. The length of such migration in search of a habitat not occupied by sables is determined by the degree of occupation of adjacent areas. With development of harvest, these migrations are not great, in other cases, sables cover a distance of 50-70 km and more.
Older sables Camans") with worn teeth lead, to a significant degree, a nomadic or seminomadic mode of life. They roam very widely
within the limits of their range in search for the most available foods.

migrations are noted above

winter, sables descend

For Kamchatka (Vershinin and Dolgorukov, 1947) seasonal all; annually, in the second half of

from the mountains into the valleys

in

connection with deepening and compaction of the snow, following

hares and ptarmigan.
Significant emigrations occur locally in years with sable food
shortages.
dies in
its

However, north Ural sable hunters believe that the sable place and does not leave the region of its inhabitation.
is

Apparently, sable behavior

geographically variable.


812
Finally, massive displacements of sable are

known

in cases of

eruption of volcanos (Kamchatka) and taiga forest fires in the dry

years that sometimes cover vast territories in Siberia. However, in
the case of large taiga forest fires, comparatively

saved by fleeing; the majority of them die from the

ently,

few sables are fire. Apparetc.,

massive migrations due to food shortage

to 15 years.

is

possible. In the

Sayans, they were noted in 1875, 1888, 1900, 1915,

approxi-

mately every 12
2.0; 2.0; 4.0

This question

is

not yet well studied.

at distances

Sables tagged in the northern Urals were caught

of

from the point of release (Raevskii, 1947). One tagged sable was caught after 8 years 20 km from the place of release (Nadeev and Timofeev, 1955). Dispersal of young sables sometimes takes place in groups (litters?), in which one sex often predominates (Durkeit,1957). The
and 4.5
length of daily
trails

km

of dispersing sables reaches 10

km

and more, the

animals moving with longer jumps than during the fattening period.

With increase
in

disperse are usually those

from which sables which are not fully exploited. They settle areas where sables were removed during the harvest period.
in

snow

depth, dispersal stops. Areas

In Glacier National Park in the

USA,

in

an experimental area

of 1,554 hectares, 85 sables [=American marten] were marked and

repeatedly trapped; 54 males and 31 females were caught 525 times,
i.e.

an average of six times each; 57.6% of the sables were caught

545

was caught 73 times during 562 days. Only 27 sables were entirely settled. Thirty individuals stayed in a place for more than three months, nine individuals from one week to three months and 47 sables were transients. Reproduction. In cage raising, the main mass of sables attain sexual maturity at the age of 2 years and 3 months. At the age of 15 months, 16% of the females become sexually mature (Starkov, 1947). In Pushkin state fur farm, 10% attain sexual maturity in the second year, 70-80% in the third year; the remaining animals in the fourth year or they do not reproduce at all (Nadeev and Timofeev, 1955). Termination of sexual activity is observed there at 13 years; individual females remain fertile until 15 years old. At 16-17 years, non-productive attempts at reproduction still occur.
not less than two times, and one male

Of

nine females older than 10 years, only one exhibited senile

changes of the sex organs, but the old females are usually less fertile. In captivity, sable live for about 15-18 years.

813

was observed between 5 and 8 years (Starkov, 1947). During the ages of 4-9 years, the average number of young in a litter is 3.5-3.6 and by 15 years, falls to 1.5. In nature, the majority of females, according to data from his-

The

least percentage of infertility

tological investigations,

(Zaleker,

1953).

become pregnant in the second year of life The percentage of pregnant females among
second year of
life in
it

Barguzin sables
to

in the

nature was from 26

33%

(Baevskii,

1956). In adult females,

ranged from 50

to

72.9% depending on nutritional conditions in the data year. In

regions of intensive exploitation, infertility

higher

(72%

as against

among females was 50%), while among young females, pregat all.

nancy was not observed

among
in

different geographical populations ranged

The percentage of pregnant females from 88 to 100%

6 different regions (Zaleker, 1953).

In

among second-year females 27.5%, among third-year 46.3%, and among older females 62.5%, and very females ranged old females 20%. Percentage of pregnancy
pregnant,

Kamchatka from 1952-1962, of 795 females 46.2% were

in

females

in

(in

average values) within the limits of 3.8% to 23.5% annually (A.A.

Vershinin). Estrus in fur-farm sables extends from 15 June to 15 August
(Starkov, 1940; Kler, 1941): from 6-25 July,

have come into estrus, and from 11-20 July 38%, the mean date lies within 16-20 July (157 cases of 789). The earliest cases are twice as frequent as late ones. In nature, spermatogenesis was histologically established for the period from 11 VI to 29 VII
(Raevskii, 1947). In August, sexual activity
is

68%

of

all

females

suppressed. Estrus

extends for 1-3 days with repeated mating, each of

which contin-

ues from 30-40 min to 2h each. After a pause of 1-10, and up to 20 days, estrus is repeated. Full sexual quiescence in sable extends

from the end of August to February (Kler, 1941). During this time, the development of the embryos, which are in the trophoblast stage,
almost stops (latent period).

At the end of February-beginning of March, following phenomena very similar to those in the esterus period (up to vaginal cornification), the placentation [implantation] of the embryos
takes place. Their subsequent development proceeds rapidly for

27-28 days
this are

(Kler,

1941).

The phenomena of

the so-called false

heat are connected with the period of placentation.

The reasons

for

not yet clear.

814

General duration of pregnancy is determined as 273-275 (Starkov, 1947) on 245-298 days (Kler, 1941); this variation is related to the variation in length of the latent period. The average
duration of pregnancy
is

riod of pregnancy in sable

273 days. The duration of the latent peis determined to a minor extent by two

factors: response to [passage of] time

in length of daylight.

and the

light factor

change

The

artificial

lengthening of the latter caused

shortening of the latent period in

Parturition takes place

50%

of experimental females

(Belyaev, 1950; Belyaev, Perel'dik and Portnova, 1951).

from 25 March-3 May, 83% of the

cases occurring in a period of 20 days (9-27 April), and

50%

in

10 days (11-20 April: Kler, 1941).

The number of young

ticular);
in

in the litter is

very tightly linked to the

degree of favorableness of external environment (nutrition in parSiberia

it

[M.

zibellina]
1

americana],
the

varies

from

to 7.

and North America [M. In the Pushkin state fur farm, the
equals 3 (Starkov, 1947); in

average number of young in a

litter

Moscow Zoo,
to

it

averages 2.8 (1.5)*. Sable in nature were

litter size

determined
546
in

have an average

of 2.5 (Kosmakov, 1936);

Barguzin sable, with the calculation of mortality before the beginning of harvest 0.7 (Z.F. Svatosh); in Kondo-Sos'vinsk sable 3.5 (Raevskii, 1947 30 cases); in sable of the northern

Urals 3.0

(2-4) (Belousov, 1914); in the Eastern

Sayan 3.0

(1-

4) (Solov'ev, 1921); in

Kamchatka 3.0

(2-4); on Bol'shoi Shantar

Island 2.0 (2-4)

(Dul'keit, 1929); in the

Koryaksk range 3-6

(Sokol'nikov, 1927); in Shitkinsk region of Irkutsk oblast

3.3

(Timofeev, 1951). The average

fertility

of Barguzin sable on farms

was determined as 0.92 and of Amur sable 1.76. The potential fertility of sable, determined by the number of corpora lutea (without calculating embryonic mortality), is variable. In individual geographical populations, the average number of corpora lutea ranged from 2.3 (Barguzin and Gornoaltaisk sables) to 4.0 (Ivdel'skii region, Sverdlovsk oblast)

and even

to 5.0

(Tyumen oblast, 1952/53) (Zaleker, 1953, 1955, 1959; Baevskii, 1956). The average number of corpora lutea changes annually depending on the general
stability of sable food; thus in the region
little

of Barguzin preserve, with

food, the average

in

number of

corpora lutea was 2.3 (1952), and with an increase

food (1953),

*Meaning of number

in

parentheses not clear

Sci. Ed.

815
increased to 3.2 (Baevskii,
this indicator

it

1956). In the Ivdel'sk region, in

1951/52
oblast

was 4.0 and in 1952/53 4.5; in Tyumen and 5.0 respectively (Zaleker, 1959). Variations in the average number of corpora lutea with age

3.1

were also noted in sables. In young female Barguzin sable in 1952, 3.0, and in 2.3; in adult females in 1952 it was 2.7 and in 1953 1953 3.4; in old females 2.7 and 4.0 respectively (Baevskii, 1956). In Kamchatka, over 10 years (1952-1962), pregnant females among the second-year females had an average of 3.9 1.08: corpora lutea per female, and in all females of this age among third-year pregnant females 4.01 and 1.86; and among

older females

4.38 and 2.71

respectively. In very old females, 3

corpora lutea were found

in

pregnant females and the average

very old females was 0.6. For the
lutea per pregnant

number of corpora

lutea in all

entire period, the average

number of corpora
to

female was 3.96, from

to 9. Calculating all females, the

average

number of corpora

lutea

was found

be only 1.93 (0.51-3.08). In

total

second-year females, corpora lutea were (from a

of 1033)

21%;

in

third-year females

30%;
A

and

in

older females

49%.

Females 2-7-year old constituting

lation, give

22%

(on average) of the poputhe latter are poten-

50%

of the potential offspring, but the second half by

i.e.,

older females, which constitute only 11%;

tially

more (by 2 times)

fertile.

definite

dependence was estabits

lished of potential fertility on the nutritional level of the female,

weight, and helminth infections (A.A. Vershinin).

Data on
it

real

fertility

of sables in nature are extremely in-

adequate for introducing corrections to potential fertility; however,

is

clear that with different age composition of the sample

populations, the results will be different. Investigations in recent

years have

shown

that in

martens with a latent period in pregnancy

[delayed implantation], ovulation takes place periodically before placentation of the embryos in pregnant females. Although these

ova are not fertilized, the number of corpora lutea increases (Neal and Harrison, 1958). Probably, this is also the case in sable and in pine marten. Apparently, the number of corpora lutea may only be
considered an indirect indicator of the potential
fertility,

since

it

may exceed
mortality.

the

number of embryos even discounting embryonic

less, blind,

Growth, development, and molt. Young sables are born helpand with membrane-covered ear openings. At that time,

they are covered with sparse light-gray down.


816

The average weight of newborn

length of 11 to 12
g,

sables

is

about 30 g with body

cm

(Starkov, 1947).

single
g.

pup may

attain

34

while in large

litters, it is

reduced to 25

Sable cubs of larger

races have a weight up to 32.8 g (with body length 11.9 cm). 547 The auditory orifices open on the 23rd-24th day; eyes on the

30th-36th day (Starkov, 1947). Incisors erupt on the 38th day, and the replacement of milk teeth by permanent ones begins at the age
of 3.5 months.

Sable cubs
Lactation ends

start

to leave

the nest at the age of 45 days.

attained 2

when cubs have

months of age. The

food begins somewhat earlier at the age of 50-52 days (ManteifeF, 1934). At one month of age the sable cub attains a weight of 260 g, 600 g (Starkov, 1947). By July (the breeding peat two months riod of the parents), they already weigh about 800 g with a body length of 39-40 cm, nearly attaining the dimensions of adult sables. With the transition to independent feeding, the growth of
utilization of animal

sables noticeably increases.

50.8%. Investigation of 1139 individuals from 1918-1930 56% were males and females 44% (Belyshev, 1950). showed the population other data (2,492), males According have been observed Differences tuted 54.4%, and females 42.6%. Kamchatka, during by regions (Nadeev and Timofeev, 1955). 1939-1962, 3,967 individuals were investigated 52.9% males were track the same place, according found, females 47.1%.
males
that
to
in

Among

195 newborn sable cubs, 49.2% were males and

fe-

consti-

In

In

to

counts, from 1950-1958,

54.2% males were found and females

In both cases
it

45.8% (A.A. Vershinin).

centage of males

is

possible that the perfirst

may be

overestimated, in the

instance caused

by selection

for harvest,

and

in the second instance, caused

by greater

activity of males.

In the sable, there

is

a spring

and an autumn molt.

Enemies, diseases, parasites, mortality, and competitors. The

young sables

few enemies among other animals. In this respect, are exposed to the greater danger. The white-tailed sea eagle, eagle owl, goshawk, yellow-throated marten, wolverine and red fox are revealed to be enemies of young sables by factual observations. In all events, direct predation on sables cannot be
sables have

considered as a factor affecting population numbers. In nature, epizootic diseases of sable are undoubtedly present,
but this question has been poorly studied.

die-off

among

sables


817

was observed in the West Siberian lowland (Raevskii, 1947), in Kamchatka (Averin, 1948), the Far East and other regions. It was
also noted locally in the 60' s because conditions of overpopulation

that

were favorable for the development of epizootics. Among diseases have been established are encephalitis, coccidiosis and skin
In the sables, 14 helminthic species have been noted (Petrov, 1941).

fungal diseases.

However,

their pathogenicity
is

is,

in the majority
Its

of cases, unknown.

Mortality of sables

quite significant.

magnitude

is

deter-

mined by many causes.

and
its

First of all, the mortality of the population

age composition depend on the intensity of exploitation and the degree of suitability of environmental conditions for the species. The magnitude of juvenile mortality (to one year) is very
considerable.

Under

the severe conditions of the

Koryaksk range

(Sokol'nikov, 1927) and Bol'shoi Shantar Island (Dul'keit, 1929),

the percentage of mortality during the first year of life

was

deter-

mined

be from 76.7 to 83.4. Just embryonic mortality lies within 8-31% (Baevskii, 1956, 1957; Belyaev and Utkin, 1960). In the absence of exploitation, the Kondosos'vinsk sable poputo

were of the following: up to one year 42.6%; 1-2 years 28.5%; 2-3 years 23.8%; above 3 years4.7%. Thus, in spite of entirely favorable ecological conditions, the
lation age groups

intensity of the mortality rate of the population here

was consid-

erable and the average age

is

not high (P.B. Yurgenson). In Barguzin

sables, the ratio of age groups observed was:

young males

moderate age

37%;

old

26%;

37%,

females were 59.4%, 31.2% and

9.4% respectively
548

(Favorskii, 1935). Mortality

found to be significantly higher than that

In 1952, in the

among females was among males.

territory of

commercial harvest on the

Barguzin

preserve,

16%.

In the following year,

young females constituted 33%, adults 50% and old, young [females] were 49.2%, adults

42%, and old

6.8%.

This appeared to be the result of intensive

year, in an area

harvesting. In that

same

vesting had long been intensive, the percentage of

where commercial haryoung was 60.7%,

at all (Baevskii,

of adults

39.3% and old females did not appear

make
it

1956). These circumstances

sity of exploitation

possible to estimate the inten-

through the age composition (Table 53; Nadeev

evident that the size of annual growth

and Timofeev, 1955).

From

all

these data

it is

in local sable

populations

may

range from 10.5 to 30-40%. In the

818
Table 53. Changes in age composition of separate sable populations in different
regions (630 individuals)

Age group
Less than
1

Vitim
year

Angara

Sayans

Kuzbass

Altai

Average

17.6%
30.9
26.7 24.8

32.0%
20.8
18.2

25.1%
25.1

14.5%
21.5
17.2

10.5%
12.5

28.5%
27.1

About 2 years
Older than 2 years
Particularly old

24.3
24.5

14.6

21.3
23.1

29.0
131

47.0

62.4

Examples studied

165

187

98

49

630

Shitkinsk region of Irkutsk oblast (Nadeev and Timofeev, 1955),

during 5 years the percentage of yearling sables changed from 23.8

36.8% (average; 31.6%). In northwestern Yakutiya, yearlings comprised 1/3 of the population, and in the Oleneksk region
to

(1950/51),

37%

(Tavrovskii, 1958).

Age

analysis of 1762 sable skulls from

Kamchatka
is

(collected

1945-1962) gave the following

results (Table 54).

The decrease
mortality

in the percent of
is

young females

explained by

their great utilization; there

information about the increased

among females from helminths (Kondrat'ev, 1957). During 1952-1962, the average percent of the juveniles in the population was 26.7%. This average changed annually for a series
of years in the following manner (A. A. Vershinin):

%
1952/5361.5 1953/5414.9 1954/55_51.9 1955/5641.2 1956/5728.6

%
1957/5814.3 1958/5981.5 1959/6043.7 1960/6115.0 1961/6250.5

For 10 years, the average percent constituted 40.3 (14.3-81.5%).

549

Kamchatka, for the same decade the size of the potential increase was obtained based on the percentage of adult females
In
Table 54. Age composition of Kamchatka sables from 1945-1962

Age
Juveniles

Total

Males

Females

27.3%
22.1

21.5%
19.8 19.3

23.0%
24.3
19.5

18-22 months
In the 3rd year

19.4

Older ages

31.2

39.4

23.5

819

and the average number of corpora lutea (A. A. Vershinin). It is given in comparison with the percent of juveniles captured during
those years (Table 55).

From

this table

may be

seen sharp fluctuations in the average

number of corpora lutea in females and hence potential fertility. However, in a series of cases (1952/53, 1958/59, 1959/60), the two latter figures are contradictory, explainable only as random selection of harvested animals as regards their age (A. A. Vershinin). In

Kamchatka, 32.5% die during the embryonic and early postembryonic periods, and in the unfavorable years
increase.

50%

of the potential

These figures

are, apparently,

lower than the actual size

of population growth.

Animals consuming nut pine "nutlets" and carnivores feeding on mouse-like rodents may be referenced as the sable's competitors. However, few of them are of substantive significance; among the consumers of nut pine "nutlets" are the Siberian chipmunk and

brown

bear, as well as forest voles [Clethrionomys] which, however, themselves serve as food for sables. Nutcracker plays a positive role; on account of the making of its stock of nut pine
"nutlets" on which the sable feeds in the snowy period of the year. Ermine, Siberian weasel, fox and other predaceous birds and animals of the taiga are considered competitors of sable in hunting

mouse-like rodents. Siberian weasel, ermine and weasel are themselves actively pursued by sable. In areas where sable settle,
the Siberian weasel disappears and
is

displaced into habitats less

suitable for the sable (Shaposhnikov, 1956),

Table 55. Potential growth of sable in Kamchatka from 1952-1962
Years Adult females. Average number
Potential

Juveniles in
the harvest,

%
1952/53 1953/54 1954/55 1955/56 1956/57
30.0 43.5
32.5

of corpora lutea

growth

1.36

40.8
36.6

61.5
14.6

0.84 2.46
1.80

80.0
48.5
28.5

51.9 41.2
28.6
14.3

27.4 36.0
43.5
22.7

0.79
3.08
2.01

1957/58 1958/59 1959/60 1960/61 1961/62

134.0
45.6
15.5

81.5

30.4
41.3
25.3

0.51

43.7
15.0

2.34 2.70
1.86

96.6

68.4
61.3

50.5

Average

33.0

40.3

820
Population dynamics. Changes in their numbers is a characterof sables. Changes brought about by deep reconstruction of

istic

natural landscapes as a result of

human

activity are irreversible:

the disappearance of coniferous taiga and the formation of the so-called cultural landscape, within the limits of which sable
usually do not survive.

To

those powerful factors influencing sable numbers, are also

applied taiga fires and the hunting harvest. Diseases and starvation are less significant, since contact between sables in the populations
is

small and the sable

is

polyphagous,

it

suffers

from food scarcity

its

infrequently, and not everywhere. In cases

when

density in-

creases, the possibility of contact grows, and serious epizootics

begin to appear
or
550

among
in

sables.

weak

(preserves), fluctuations in sable

Where human activity is excluded number are insignificant

combination with and as a consequence of, in some seasons, an accumulation of favorable or unfavorable environmental factors. In American marten, a certain cyclicity was observed in fluctuations of their numbers. Years of high number

and are determined

are repeated,

on average, every 10 years, with deviations

in the

range of 9-11 years.

Kamchatka, voles are of decisive importance as sable food (see above). Their mass reproduction there has a three-year cycle;
In
i.e.

similar to the so-called "Scandinavian cycle."

As noted above,

the level of fertility depends on the age structure of the population,

caused by lower
duction
tains a

fertility

of young females. With intensive reprois

when

the

main food
is

plentiful, the sable population atin the

younger average age and therefore,

following year,

breeding intensivity

reduced. In the third year, the age structure

becomes even
and
it

less favorable for productivity of the population,

falls to

minimal increase. In the fourth year, the relative

proportion* of the oldest age group increases, and the population growth again increases. Thus, the numerical dynamics of the population
is

determined by the
in

availability of the
in

mental factors which determine changes

main food: 1) environabundance of voles, and 2)

fertility.

by changes

population structure which change the level of

In those parts of the range where a considerable variety of alternate foods are present and the sable does not depend on the abundance of 1-2 foods (usually interrelated, for example, voles and nut pine "nutlets"), its numbers are subjected to smoother
*Iii

Russian original,

literally specific

weight

Sci. Ed.

821
fluctuations, depending on a combination of different factors and the

cycle approaches 9-11-year interval.

If,

in

Kamchatka, the numerical

is

fluctuation attains an interval of 6 [years], in such regions with a

prolonged cycle,

it is

2-3 times lower;

i.e.

the population

more

stable.

In the Glacier National Park (Montana,

USA),
in

the process of

numerical change in concrete local populations

an area of 1,554

ha was studied by means of tagging and recapturing animals (195254). At the end of the summer of 1953, population growth was observed. After that, from September 1953 to December 1954, of 27 resident sables, nine were killed (30%). All the progeny of 1954 also migrated beyond the boundaries of the experimental territory. Density fell from 17.6 to 7.0 per 1000 ha; i.e. by 2-4
times. Five females died

(18.5% of the population). Decline

in the

weight of trapped females was noted, and dead animals showed

signs of emaciation and shock disease (Selye's syndrome). There

was

first

overpopulation and exhaustion of food resources owing

to reduction in

numbers of the principal food

mouse-like rodents.

Predominant mortality of females was explained by increased loss of energy (estrus, pregnancy, parturition, lactation) due to insufficient food. Moreover, in those places where females fed, their litters fed with them, and this accelerated the exhaustion of food stocks (Newby and Hawley, 1954; Hawley, 1957).
Field characteristics. Under natural conditions, most often only
traces of sable activity

must be dealt with. The animal

it is

itself

may

be met with rarely and accidentally;

resembles a small
cat,

easier to succeed with the

help of a good laika dog. The sable, driven up a tree, somewhat

but differs in
its

fluffier fur, small sharp

muzzle and

relatively short fluffy

tail. It is

very similar to the pine

marten, but usually

may be

distinguished by the lighter color of the

tail.

head and somewhat shorter

tiated

The male can be

easily differen-

from the female by

its

larger dimensions and muscular, burly

shape; the females are thinner and more elegant. Tracks of sable are
very rarely found in the snow-free period of the year, in the form of
single prints.

One may
It is

often find sable scat on taiga paths, on stumps

and

fallen logs.
in diameter,

sausage-like in

form with blunt ends, about

it

1.5

cm

and a length of about 6-7.5 cm. In

are usually found

hair

and small bones of voles,

shells of nut pine "nutlets", fruit

and

insect remains.
551

The

scat gives off a

musky

odor.

Footprints of sable are broad, and disproportionately large

compared

to the animal's

body

size.

The

large footprints of a sable


822
track are often not differentiated

from those of red

fox.

The

feet

of sable are not held in a clump, distinguishing them from red fox.
In a clearly printed sable track, the claws are not obvious; foot

pads are not always visible, but are more frequently seen in summer. On packed snow, the average dimensions of the prints are
equal to 4 x 6.5

cm

(from 5 x 7 to 6 x 10 cm). Doubled prints of

front and hind feet are usually large

from 5x9

to

8x15

cm. In

large leaps on soft

snow

the prints of both feet are united in one

On packed snow, sable form depressions 3-5 cm, and on soft snow 10-15 cm. In addition to paired prints, especially on packed snow, sable often trot. Then, the arrangement of the tracks is very similar to those of hare. Sable walks "cleanly" without leaving drag marks.
depression of dimensions 10 x 25 cm.
tracks

In the Siberian taiga, tracks of sable are similar only to the

tracks of Siberian weasel, but only inexperienced hunters might

confuse

them. The tracks of the Siberian weasel are not located

parallel to

at the heels; this is less

kidas.

somewhat overlap each other marked in the tracks of pine marten and When jumping from a tree, the pine marten does not leave
each other, as
in sable, but

the

tail

print

on the snow, but the sable does. Passing under the

its

crown of

a tree, the sable greatly shortens

leaps, but the pine

marten continues walking regularly. Moreover, sable never stays twice in one shelter, while the kidas does; sable go about widely under the snow completely covered with it. Pine marten and kidas
can
sit it

out under the

snow

to the very last

extreme (Raevskii,

1947; Yurgenson, 1948).

In the Urals, after a snowfall, sable begins to

move

after

2-3

days,

when the snow has more than 3-5 cm.

settled

and when the animal sinks

in not

Usually, the sable does not run the whole day with one gait.

upon snow density. In soft snow, its feet more widely. In soft snow, length of the leap does not exceed 30 cm; in dense snow 45-75 cm. Separate large leaps reach 2 m; such leaps do not occur in more than 2-3 successive jumps. The average length of a leap in the montane taiga of the northern Urals is equal to 40-45 cm, ranging from 70 to 80 cm (Kondratov, 1953).

The length of

the leap depends

leaps are shorter and the sable places

In the taiga of the eastern Sayan, the length of sable leaps

ranges from 40 to 80 cm. The width of the paired tracks usually ranges from 9 to 14

cm

and

that of

one track

from 4.0

to 5.5

cm.


823

The

sable foot sinks into the

snow

for

5-10 cm. Weight loading

is

equal to 12-14 g per 1 cm^ (Dul'keit, 1957), in the northern Urals, even with a body weight of 1200 g, it ranges from 9.5 to 11.7 g (Kondratov, 1953). The average length of a leap in a male Sayan
sable in soft

snow

is

equal to 51.5 cm, of a female

37.5 cm; the

(9.4-11.9), of

paired prints of a track in females average 11.0

a

cm

male 13.3 cm
In the

(12.0-14.6).
gait,

most typical

which make paired

is

prints, the poste-

rior pair of feet lie in the prints of the anterior pair.

is

One

of the feet

usually displaced forward, and there

the right and left feet.

feet

triple print, in

no regular alternation of which one of the posterior

is

does not

lie in
is

the print of the anterior foot,

infrequently

observed and

usually noticed
It

when

there

is little

snow, when the

feet separately

sable feels firm support.

(as in red fox).

rarely places all of

its

The speed of movement of sable is 7-8 km/hour, and in one minute it makes 200 leaps, of 60-70 cm each. Walking, the sable ambushes its prey; it steps, it listens, considers etc. Individual sables can easily be distinguished from one another by the length of leap, the width of feet placement and depth of sinking into the snow, since these values depend on the weight and size of the sable. In order to travel 1 km, the male makes on average 1900 leaps, and the female 2400, but in soft snow 2500 and 3300 respectively. Consequently for the same distance, the female requires more energy and therefore, its daily trail is always shorter. The length of the trail is shortened with an accu-

mulation of soft snow, when leap length decreases and sinking of the animal into the snow increases (Dul'keit, 1957) (P.Yu.).
552

Practical Significance

The sable
same value

is

the

most valuable fur-bearing animal

geographical regions and even

in the
is

USSR
same

although, as a commodity, the quality of the skins

in different

not of the
in the

locality (see above, "Description" section).

Excessive, often rapacious, exploitation of sable in the past

exhausted the stocks of

this animal. Its

range was also strongly

reduced. In order to restore their number and range, a series of preserves were organized, their exploitation was prohibited for a

long time and broad encompassing measures for artificial reintroduction of sable were undertaken in regions where it had been

824
extirpated.

As

a result, the

by

it

increased considerably. The

number of sable and the area inhabited number of sable attained such

heights that in

some

parts of the range a surplus of sable devel-

oped with

a series of negative consequences. Captive breeding of

sables developed widely and attained significant success.

At the present time, within the boundaries of the USSR, about of the sables caught each year come from Krasnoyarsk Territory, about 20% from Irkutsk district, about 16% from Khabarovsk Territory. With all of this, at the present time, not all

33%

the areas in the Siberian taiga suitable for their occupation have

been taken up by sables. At the present time, the tendency

wards increase
there stands the
in the

is

to-

number of

light-colored sables. Therefore,

selection of sable with the entire

problem of devaluation in the direction of natural improvement in the quality of the

skins on a large scale.

A
lation

very important measure for management of the sable popuis

to determine correctly a guide to the

amount of
it.

utilization

(the percentage of the admissible catch)

and maintain

For a very

high density, the catch should be accelerated. The example of

Kamchatka can
sand sables
in a

illustrate this situation: a catch of

7.5-8.0 thou-

season does not lead to a decrease

in their

number,
it

but a catch of even 8.5-9.0 gives this effect. In Kamchatka,

is

admissible to take

27%

of the autumn pre-harvest number (A. A.

Vershinin).

In

other

places 25% and 30% (Baturin, 1930;

It

Vershinin and Dolgorukov, 1947).

is

important to change the

size of the catch in relation to the condition of their numbers: for

Kamchatka the limits are between 15% to 65% (A. A. Vershinin). The main methods of sable capture are hunting with guns, with
the aid of laika dogs, harvest by trapping, and the application of
baits. Different

wooden
is

deadfalls, snares, and live-traps of the

box

type are also used. The hunting of sable by baits helps to obtain

them

alive,

which

important for the purpose of reintroduction,

selection, banding, etc.

The sable harvest

is

regulated by the issu-

ance of special licences.

Measures for the effective protection of sable should include: maintenance of rules and norms of licenced harvest; 2) prohibition of selective destruction (which is possible by baits of
1) strict

high quality sables through establishing "average" prices for each

fur type. Otherwise, the hunter, having a sable hunting licence,

will

always

try

to

take only sables of the highest sort, thus


825
increasing the low quality sables in nature; 3) banning overharvest
in core habitat (as regards quantity

and quality) throughout the

enitre range, as well as other measures.

It

is

desirable to prohibit periodically the harvest, even with

licences, after seasons unfavorable for sables (famine, forest fires,

diseases and others) and to organise supplemental feeding in the

winters of famine.

It is

also necessary to put into practice a harvest

with planned selection of sables in nature through issuance of

special licences for trapping
ity sables

and shooting of only the lowest qual("mekhovaya" and in some cases, "vorotovaya").

553

The greatest efforts should be directed towards the development of excess populations in unharvested and poorly harvested
regions, and towards the prohibition of overharvesting in areas

lying close to the inhabited areas, and those

easier to develop (P.Yu.).

more convenient and

FOREST MARTEN26*
Martes (Maries) martes Linnaeus, 1758
1758. Mustela martes. Linnaeus. Syst. Nat. Ed. X,
1, p.

46. Upsala,

southern Sweden.
1816. Mustela sylvestris. Oken. Lehrb. Natgesch.,
stitute for
2, p.

1029. Sub-

martes.
1, p.

1820. Mustela sylvatica. Nilsson. Scand. Fauna. Dagg. Djur.,

41.

Renaming of martes.
5, p.

1827. Martes vulgaris. Griffith. Cuvier's Anim. Kingd.,

123.

Renaming of martes.
1865. Martes abietum. Gray. Proc. Zool. Soc. London, p. 104.

1926. Martes martes lorenzi. Ognev. Uchen. Zap. Sev.-Kavk. Inst.

Kraevedeniya,

1, p.

47. Storozhevaya,

Kuban
p.

district.

1926. Martes martes ruthena. Ognev. Ibidem,

49. Dmitrovsk,

Moscow

province.

1941. M.(artes) m.{artes) ruthena borealis. Kusnetzov. Tr. Mosk.

Zootekhn.
district.

in-ta, 1, p. 126.

Nomen

Shenkursk region, Arkhangel'sk praeoccupatum Mustela flavigula borealis

Radde, 1862.
^*Also called "zheltodushka" [yellow-breasted] and "kunitsa

myagkaya" [smooth

marten] by furriers.
* literally, "forest

marten"

Sci. Ed.

826
1941. Martes martes uralensis. Kusnetzov. Ibidem,
p.

126. Vic.

Miass, southern Urals.

1947. Martes martes sabaneevi.

Jurgenson. (Yurgenson). Tr.

Pechoro-Ilychsk preserve,

Pechoro-Ilychsk. zapov.,

5, p. 179.

upper Pechora (V.H.).

Diagnosis
Tail length with terminal hairs

more than half

that of body.

Color

chest, usually varying intensities of yellow color,

monotone, sharply outlined patch of various forms on throat and sometimes white or almost white, and as a rule, not forming two strips extending
to

backwards
sable.

base of forelegs. Head not lighter than back.

Bony
in

auditory bulla relatively shorter and

more widely separated than

Inner half of upper molar considerably larger than outer

(V.H.).

Description^^
general appearance, the pine marten in winter pelage

In

its

is

very

and longer tailed. Tail length with terminal hairs is more than half the body length in the animal "in the flesh", the tail extends behind the
similar to the sable, but looks
larger, coarser

somewhat

stretched hind legs usually for

more than
is

a quarter of
tail

its

length. In

contrast to the sable, the terminal hairs of the

blunt, but a tapering tail tip. This tip

do not form a

554

sometimes strongly elongated (majority of cases), typical of marten, sometimes tapering to an obtuse cone-shape, and sometimes (minority of cases) has the blunt "sable-like" form (Urals; Pavlinin, 1959). Length of the terminal hairs, in typical cases may reach 100 mm. Foot pads are less furry, and appear to be less broad. Ears, in sable, are also broad at the base, but shorter and more rounded at the tips, from which the whole head looks relatively smaller than in the sable, and more elongated, with a longer facial portion. Feet are covered below
with dense and quite coarse hairs, somewhat less developed, however, than in sable, the sole and digital cushions are completely

covered with hairs, but the ends of the claws protrude from the

fur.

"Some

additional data, characterizing the pine marten, are given in the descrip-

tion of the stone marten.

827
In

summer

coat, the

marten

is

distinguished from the sable by

the

same

characteristics, but appears

even coarser and more longits

tailed,

with less broad feet and not so big-eared. In

is

manner of

standing and moving the marten

differs considerably

entirely similar to sable, but

from the stone marten, which mostly "crawls"

is

and rarely proceeds by jumping. This

height of the animal
tutes about 15

connected, in part, by the

pine marten, this consti-

at its shoulders: in the

cm

(Schmidt, 1943) and more than in the stone marten.

Fig.

208. Pine marten, or zheltodushka [yellow-breast], Martes (Martes) martes L.

Sketch by A.N. Komarov.

828

but
it

The pelage of marten is similar in structure to that of sable, is somewhat rougher and less dense and fluffy. Nevertheless,

555

is one of the smoothest and most delicate furs and one of the most valuable. Compared to winter, the summer fur is short, coarse and relatively thin. In summer there are 4-5 underhairs for each guard hair. Length of the guard hairs is 23-27 mm, with a thickness of 100-150 microns, length of underfur is 11-12 mm, with thickness of 15-16 microns. The corresponding figures for the winter fur: guard hair, 36-38 mm and 75-90 microns, and underfur,

22-22.5
ter hairs

mm and

14.5-15 microns (Pavlova, 1951). Thus, the win-

although longer, are relatively

much
is

thinner. In the

sum-

mer coat of marten,

the sole and finger cushions as well as the

tail

claws are easily seen. In winter, the

much

fluffier than in

summer and with longer terminal hairs. The general color of the winter fur

is

brownish-reddish or

reddish-chestnut with a strong straw-gray wash, depending upon the general tone of the underfur. This wash is particularly devel-

oped on the sides, where the underfur is not so completely covered by dark guard hairs, as on the back. The back is darker than the sides, as are also the middle of the abdomen and the inguinal region, but lighter than the middle of the back. The top of the head is dark, no lighter than the back. The upper edges of the ears and
their inner sides are whitish-straw, the reverse side of the ear

dark

brownish. The feet are blacker and darker brown than the back. The basal part of the tail is colored like the back, and its tip is
darker.

light patch with sharp outlines is located

on the throat and

the lower surface of the neck. In a typical case, the patch occupies the throat and the lower surface of the neck, without extending to
its

sides but projecting

backwards as an extension onto the chest

between the forelegs. Its borders are usually sharply outlined, but they are sometimes diluted on the neck. In general, the form and dimensions of the patch are variable. Usually it develops in a more or less typical shape, but sometimes is rounded (backward extension
is

absent) or greatly reduced. In

some

instances,
into

it is

divided

into separate parts and, rarely,

may be reduced

some small

white marks, or disappear completely. Sometimes there is a dark spot in the middle of the light field. Sometimes, on the contrary,
the patch
is

enlarged and covers not only the throat, but the ante-

rior part of the chest

and the upper part of the forelimbs. In some

829
cases, though
is

it

very rare, there are two extensions towards

forelimbs, as in the stone marten.

As an

exception, the posterior

extension of the patch may, on passing between the forelimbs,

extend quite far along the abdomen. There are some geographical tendencies in changes in size (not form) of patch (see below).
the general color type is retained, its tone and undergo considerable individual fluctuation. One may distinguish four main color types of martens which are accepted in
intensity

Though

the fur industry (furriers terminology).

Dark blue
Blue

fur dark-chestnut without reddish tints, underfur

tip.

bluish-gray at the base and light-gray at the

Dark sandy more distinct on

color at hair

pelage
tips.

lighter, chestnut colored,

underfur gray.

brownish ("dark-sandy") with reddish tints, the sides. Underfur gray at base and light-sandy
fur
tint.

Sandy

pelage light-brownish ("sandy") with light-yellow

at

Underfur gray

base and yellowish

at the tip

(Kuznetsov, 1952).

Sometimes, skins have "grizzled" highlights scattered white guard hairs; however, these grizzled ones are much less developed than in sable and are rarely encountered. Under various conditions, "dark blue" martens are considered the most valuable. The same tendencies are also noticeable in
geographic variations in marten
fur,

however, they are quite weak

and much weaker than

itself is less

in sable

and the phenomenon of parallelism

marked.

556

the throat patch is, as a rule, yellow of varying changing from slightly ocherous to bright intensive ocherous, almost orange. Near-white and white patches occur. There are several geographic tendencies in the intensity of patch color. Among martens of the European part of the USSR, the most southern (Kuban) martens have the brightest orange patch; northintensity,

The color of

ward, intensity of
picture
is

its

color decreases, and

it

lightens.

The same

also observed in the Urals

from the north

to the south-

ern extreme of the range. Along with this, the general size of the

patch changes with

some

regularity. In the Urals, the

number of

martens with a large throat patch increases from north to south. In

Bashkiriya, martens with small patches are relatively very few. In
a

few instances (more often

in the northern Urals), the throat patch

may completely

disappear. Such individuals in several sections of

the range in the Urals constitute 1-3.2%, sometimes even to

7.1%

830
(Kuznetsov, 1952; Pavlinin, 1959). White or near-white patches
for stone marten.

sometimes found, or having a form, similar to the form typical Such individuals are very rare, and only one among several hundreds are met with in the Urals, one among
are

900 (Pavlinin, 1959)^8. The density, luxuriance and fineness of the pelage i.e. fur quality, undergo marked geographical changes which are, however, much less than in sable (see section "Geographic variation"). Individual and geographic variation in marten are on the whole, much
less than in sable.

Geographical localization of types of individual

is

variation so characteristic for sable

absent in marten, or

it

is

more weakly manifested.

Summer

fur

is

characterized by yellowish-brown color of the

underfur and dark-brown color of the guard hairs. The relatively

strong development of the guard hairs, with dark underfur, confers
a general

dark-brown color

to the animal,
its tip,

than in winter. than the trunk

The

tail,

particularly

which is much darker and the feet are darker

is

blackish-brown. The
is

throat patch

usually brighter

than that in winter marten. Individual variation in color of

fur of martens

summer

much

less than in winter.

in color

There are no sexual and age differences

of the pelage (fur quality).
that is not distinguishable

and character

Young
is

in the first

autumn put on a coat

adults.

from the winter coat of

The
It

skull of pine

marten

very similar to the skull of sable.

from it by the not so elongated brain case and by its more widely separated bony auditory bullae the distance between them in the middle part of their length is more than half the length of the bulla, measured from its anterior edge to the posterior point of the paroccipital process. The mastoid processes slightly extend outwards behind the edge of the auditory openings. The interpterygoid cavity is somewhat wider than in sable. The
differs

shorter and

dental formula

is

as in sable, but the inner blade of the upper

molar

is

usually

somewhat
is

larger and the difference

between the

inner and outer blades

greater.

Sexual differences are expressed in the smaller general measurements of the female skull, the weaker development of

^'Particularly such individuals, apparently,

were given as the reason for the ap-

pearance from time to time of confirmations about hybridization between the stone

and pine martens, originating mainly from fanciers but penetrating later into scientific literature, which in actuality does not occur (see below, chapter on stone marten).

831

557

Fig. 209. Skull of pine marten,

Martes (Martes) martes L.

832
protuberances, crests,
etc.,

and on the whole,

lighter structure of the skull. In the martens of

the condylobasal length of the

male

skull

is

in the somewhat Volga-Kama basin, more than 78 mm and

the

zygomatic

width

is

more
76

than

45

mm;
42

in

females,
(Aspisov,

correspondingly
1959).

less than

mm

and

less than

mm

Age changes
is

in the skull are significant and, in general,

correspond morphologically to those described for sable (age conformity

not established).
the
tail
is

The skeleton of

composed of 15-22

vertebrae,

i.e.

558

on average, several more than in sable; Caucasian martens have 18-21 caudal vertebrae; Pechora 17-20 (175%, 1840%, 19 25%, 20 30%; Yurgenson, 1947). Male martens of the middle and southern Urals and Bashkiriya have 15-20 (n 100), with 15

found very rarely, 17 also; most often, individual variation lies within 16-20 or 18-20, 17.9-18.7 on average. In females (n 84) the number of caudal vertebrae is 16-20, with 16-17 often found,
the average lying between 17.6-18.9 (Pavlinin, 1959).
cles,

The

clavi-

which are rudimentary in all species of the genus, are articulated neither with the sternum nor with the scapula and lie in the muscles; they are weakly curved, having their basal part shorter, the distal part longer, flattened, thinning distally and tapering at
their ends; their length is about 13

mm.

Both projections at the distal end of the os penis unite with each other and form a closed ring. Only in young animals they are
often separated, as in sable (Yurgenson, 1947).
In dimensions, the pine marten correspond
sable, usually averaging
less than the

more or
latter.

less with

somewhat body length is 380-580 mm, tail length of hindfoot, 62-93 mm, and ear height, 40-45 mm. Sexual differences in dimensions are quite marked (Table 56). Average values in the different parts of the region mentioned 422-439; differ in the following manner: body length of males
species,

Within the length, 170-260 mm,

females, 387-405 mm; tail length of males, 195-207; females, 185-195 mm; hindfoot length of males, 81.5-85.8; females, 73.9-

78.0

mm;

ear height of males, 45-47; females, 44.5

mm;
skin.

weight of

males, 795-850 gm, females, 542-620

skin constitutes

gm^^ Weight of
male
it

the female

68.4% of

the weight of the

skin,
i.e.

"Weight of carcass given without

weight of a freshly skinned hide
(Pavlinin, 1959).
It

is

actually

somewhat

larger.
to

The

in the

same region comprised from 96

159

gm

is

possible that

tation of the relative

(body length?), are also dimensions of the different sexes.

some other measurements provided by the author not standard, but are of the same type and give a represen-

833
Table 56. Dimensions of male and female martens of the middle and southern Urals and Bashkiriya (from material of Pavlinin, 1959)

Dimension

Males, n

Females, n

Tail length

(mm) (mm) Hindfoot length (mm) Ear height (mm)

Body
length

93 104 84

Weight" (gm)

405473 185228 7893 4547 6 59 6701050

102

100 78 4 49

365420 170210

7084 4346 484850

Condylobasal length of male

71.1-83.0
38.2-49.2
31.8

skull, 72.1-88.0 mm; females, mm; zygomatic width of males, 41.4-53.8 mm; females, mm; skull height in males, 27.4-34.9 mm; females, 27.0is

mm

(from material of Kuznetsov, 1941). Weight

580-1800
is

gm.
In adult martens (older than 2 years), length of the os penis

38.9-45.7

mm,

M 42.3

mm;

weight, 0.20 to 0.35 gm,

0.29 gm.

In young, length is 35.1-38.3 mm, 0.16 gm (Popov, 1943; Yurgenson, 1947; Aspisov, 1959). The preputial part of the male copulatory organ is short about 26 mm long (Shtreili, 1932). In martens of the Urals and Bashkiriya, length of the intestine of males is 157-245 cm, females, 140-197 cm; heart weight of males, 6.3-10.6 gm, females, 5.4-8.4 gm. The heart index is 8.5-

9.2%o, but
is

may be from

4.3 to 11.5.

Weight of

the liver of males

18.3-46.9 gm, females, 14.5-35.2 gm. The percentage relative

to the

body weight

constitutes

27.0-33.8%

in different populations

(Pavlinin, 1959)30.
559

where sable and marten occur together Pri-Urals and beyond the Urals, hybrids are formed between them, known as kidus or kidas. It is agreed that kidas are fertile, both between themselves and in back-crossing. Only experimentally (on farms) was there established a restriction on kidas fertility females are fertile in backcrossing both with sable and with marten; in this combination, males were shown to be infertile. Fertility of kidas among
Kidas.3' In those places

and particularly

in the Urals, in the

'"The given figures,

and assigned
features in

some at least, were apparently taken from frozen carcasses weight of the body without hide. See the data on other internal the present work.
to the

"There are very many reports on kidas and separate notes on it in the literature. Yurgenson (1947) discussed this question most completely on the basis of good material. The information provided here, if not attributed, was taken from this study.

834
themselves was not established. There were not, apparently, sufficiently extensive experiments of this kind. This complex hybridization mentioned above leads to a
situation that

on one

level individuals with

intermediate characteristics
type, and also those in

more or less defined more similar to martens "marten-like" which the sable features prevail ("mixed

pasture sable" as they are referred to Pechora hunters), and finally,

some which possess

novelties of fur quality, proportions, dimen-

sions (heterosis), etc. Certain individuals do not exhibit external

features of hybridization and their actual nature can only be deter-

mined by special investigations (skull, skeleton). Because of this heterogeneity of kidas populations, their characterization is extremely difficult and cannot be given in the form of a diagnostic key. "The range of individual variation (of kidas,
V.H.)
is

extremely great. For almost every feature

we may

find

deviations towards one of their original parental species up to

complete identity, as well as intermediate characteristics. Finally, there are also localized and sharp modifications of deviation, as well as phenomena of innovation and heterosis. All these are
encountered
in multiple different

combinations. Therefore, the iden-

tification of kidas is only possible

on the basis of a combination

of characteristics, or by simultaneous unification of features of

both original species" (Yurgenson, 1947). Externally, the typical kidas of the upper Pechora

[river] are

usually characterized by the whitish color of the head, as in sable, the tail, short as compared to marten, but longer than in sable, and

by pelage that is usually dark or black in general color and denser, longer and more luxuriant than in marten. The fur, however, is coarser than in sable, with denser and thicker guard hairs, and without such a lustrous and silky appearance. In some cases, the fur closely corresponds to one or the other of the initial species, and its hybrid nature is revealed in other features (more often in length and fluffiness of the tail).
Particularly characteristic
is

the usually well demarcated, bright

colored throat patch. In some cases, coarse fur occurs without the silky appearance and luster, with dense and long guard hairs differing

from the

fur of both initial species. In this case, the animals

usually manifest particularly large size, a massive, rough appear-

ance and a heavy angular head. Concerning the general color type in a series of kidas from the Pechoro-Ilych reserve, 54% have the

835 marten type and

46%

the sable. Concerning

tail

length,

40%

are

closer to marten and

60%

to sable.
size;

Kidas of the same reserve are also of different

average they are smaller than sable (which
is

on the

marten in the Pechora), but larger than marten. Individual animals are larger than the largest martens and are nearly of the same size as
larger than
560 are as follows:

The dimensions of kidas (24 males, 10 females) body length of males 390-M 452-485 mm, females 365-M 406-450 mm; tail length of males 170-M 187240 mm, females 1 10-M167-180 mm; hind-foot length of males 75-M 86-98 mm, females 70-M 73-77 mm. Length of
the largest sable. ^^

OS penis

36.7-M 40.6-44.7 mm.

Weight of males 600-M 754-980 gm, females 410-M 551-680 gm.

In relative length of tail, kidas occupies an intermediate posi-

to a

between sable and marten. In sable of all ages, this is equal ratio of 2.7 (females, 9) and 2.9 (males, 14); in marten, it is 2.0 both in females (213) and males (251); and in kidas, 2.5 (females 6) and 2.4 (males, 6) (Yazan, 1962). Condylobasal length of skull 74.8-M 81.5-88.2 mm; zygomatic
tion

mean

width 52.1-M 55.6-58.1

(10 spec).
than

mm;

skull height

36.1-M 38.5-42.8

mm

The structure of the os penis in the majority of cases (more 90%) is that typical for marten (closed ring at the end).
Craniological features of kidas also delineate a very compli-

cated picture. Thus, out of the 40 skull measurements of Pechora

kidas,

and

35% have an intermediate characteristic between martens sables. On average, 17.5% of measurements are similar to

those of sable and 47.5% are like those of martens. For example,

zygomatic width, length of nasal bones, width of cranium and others

are sable-like; length of

auditory capsule, postorbital constriction,

skull height, etc. are marten-like.

The
41.6%
559

individuals

molar structures among is as follows: 41.6% of kidas have "sable-like" teeth, "marten-like" type, and 16.8% intermediate characters.
distribution of different upper

"According

to other data

larger than both martens (463),

(Yazan, 1962), kidas of Pechora-Ilych reserve (14) are and sables (23) of the same place. The discrepancy in

the data of Yurgenson (1947) is explained by the fact that the latter compared Pechora ^^Q martens and kidas with Kondosos'vin, i.e. the Trans-Ural sables, which are larger than the Pechora. Absolute figures presented by Yazan (1962) could not be used because
the

means of obtaining them

fur, or

are not clear (fresh carcasses with fur, frozen carcasses

with

frozen skinned carcasses?).


836

marten-like postorbital constriction

is

present in 8.3% of kidas,

It

sable-like

33.3%, and

transitional type

58.4%.

was noted

that

in individual

kidas more features of one or the other species occur.

In practice, very multiform combinations are

met with

in

kidases: os penis of the marten type, postorbital constriction and

pelage of the sable type,

features,

tail

as in pine marten; in craniological

more sable

characters, etc. Clear intermediate forms are

sharply distinguished, such as light Ural sable with long martenlike tail.

In different places, evidently in connection with the correla-

between initial species, the kidas population deviates more to one or the other side. At the sources of the Pechora, the marten type is more common, beyond the Ural, the sable type. The percentage of kidas in the population may be very high 58.8%, kidas 29.6% and for example, on the Pechora: marten 11.6% or 80%, 7%, 13% respectively. When marten and sable kidas only were hunted (sable was prohibited), at the sources of the Pechora river and along the Ilych, the ratio of marten to kidas in different places was 85 and 15%, 79 and 21%, 72 and 28% and
tion

even 65 and
561

35%"

(V.H.).

Systematic Position

See sections on sable and stone marten.

Geographic Distribution
In Europe, western Siberia, Caucasus, Asia

Minor and

Iran.

Geographic Range

in the Soviet

Union
It is

This constitutes about half of the species range.

located in the

western, and in part the southwestern, sections of the

USSR. Within

"After completion of the manuscript of this volume, the work of Pavlinin ("Tobol'sk sable", Sverdlovsk, 1963), in which, contrary to the accepted idea, the existence of kidas is denied. According to the conviction of the author, individuals considered as hybrids are merely individual variants of sable or marten. Those general considerations that are reported in connection with the existence of these inter-specific hybrids (absorption and displacement of one species by the other,
appeared
the formation of races of, hybrid origin, etc.) are also rejected.

At the same time, the author assumes

exceptional conditions" (page

that kidas in nature all "appear under

105), providing in substance the

same reasons

that

other authors have argued for the appearance of kidas. This question requires further
study.


837
the limits of our country, the range

is

divided into two parts

European-Siberian and Caucasian, which are completely separated

from each other by steppes, but are united farther to the southwest. The northern border of the range in the west starts on the Kola
Peninsula, at the state frontier at the latitude of Kola, or a
the north (the most northern point of occurrence in the
little to

USSR), and

passes to the southeast along the forest edge (Pleske, 1886) ex-

tending towards the White Sea along the southeastern coast of the

Kola Peninsula. Farther on, the border is formed by the White Sea which the range almost everywhere, extends except, apparently, to a section northwest of the Kuloi river mouth. On this part of the coast, the border crosses the lowermost Mezen' and Kuloi rivers (Zhitkov, 1904) eastward including the upper course of the Semzha (flows into Mezen' Gulf a little north of the Arctic circle) and passes through the southern part of the forest-tundra in the basins of the Peza (right tributary of the Mezen') and Tsil'ma (left tributary of the lower Pechora) rivers. East of the Pechora, the borderline passes approximately along 67" N. lat. (V.Ya.
coast, to

Parovshchikov).

Beyond

this line

long transgressions occur

the

sula to lokan'ga on the [White] Sea coast, 50

km

Kola Peninfrom the forest

border (Smirnov, 1901) and

Kiya station, on the western shore of the Kanin, approximately 200 km north of the place of permanent occurience at the mouth of the Semzha (V.Ya. Parovshchikov). According to 30' s data, the border of the range in the Urals and western Siberia inclusively took the following form. It crossed the Urals somewhere at the region of 65 N. lat. (Ognev, 1931; Adlerberg, 1935), and along the eastern slope of the Urals, turned abruptly to the south, not extending onto the plain, and not crossat

ing to the east of the Sos'va river. Approximately, at 61 N.

lat.

the

borderline sharply turned back to the east, and at the watershed of

the Tapsyi (Sos'va basin) and Tavda (Pelym), passed onto the West Siberian plains. Passing eastward, it crossed the sources of the Konda river, ascending along the left tributary of the Ob'
the Endyr' river

and

at

about 62 N.

lat.

crossed the Ob' river.

Beyond
where
it

the Ob', the range occupied the region of the lower

Nazym

reached 62 N.

lat.

Farther on, the border extended southward along the Irtysh

river to 60 N.
lat.

and again sharply turned back

to the east.

838

Following
it

this parallel, in general, or a little farther to the north,

nearly reached the Ob' (Nizhne Vartovsk yurts at the

mouth of

the Vakh).

From

this place, the borderline directed itself south-

wards to Vasyugan at the mouth of the Nyurol'ka (Nyarel'ka) and, forming its southern border, it turned around towards the southwest to the sources of the Dem'yanka (Skalon and Raevskii, 1940).
Thence, moving
in the

same

direction,

lower Ishim and, proceeding a little Golyshmanov and Yalutorovska regions,

crossed Irtysh and the northward to 56 N. lat., through

it

it

passed on to the Urals

it

through Sysert' to Nyazepetrosvk. Thence,

directed itself southits

ward

to include the Ural [river]

extending into

valley to reach

Chkalov.

An
the

isolated section of occurrence, analogous to those found in

European part of the USSR (see below), is located in the Sanarsk pine forest and in the steppe west of Troitsk (Shvarts, Pavlinin and
Danilov, 1951).
Therefore, the range of the marten in Siberia itself forms a
triangle, the

apex of which

is

directed to the east.

lat.

It

has as

its lat.,

base

the Urals between approximately 65" N.

and 56 N.

and

nearly reaches the middle Ob'*.

563

Information

at the

beginning of the 50' s (Laptev, 1956) show

a range of larger dimensions. If for several places this change

undoubtedly depends on more accurate determination of the actual

situation, then, all the

same,

it

must be considered

to

occur mainly

because of the broadening of the range, which is connected with the general increase in marten numbers and their spread to the
east.

Along

the left

bank of the Ob', marten are observed

at

in the

Shuryshkar region,

approximately 66 N.

lat.,

i.e.,

only a bit

more

to the south than

it

on the right bank of the Pechora. Apparalso crosses the Ural [mountains].

ently, at this latitude,

Berezov on the Ob' and, going eastward, encompasses the basin of the Kazym the Khetu and Sorum rivers, the Trom-Yogan basin (TromAgan, right tributary of the Ob'), and the Vakh basin (Sabun and Korlika rivers in their upper reaches). The marten was also noted at Nizhne Vartovsk at the mouth of Vakh and on the Ob' at Lukashkin above the mouth of the Vakh. The range includes the basin of Bolshoi Pasol river a tributary of the Ob', and the basin
Farther, the border descends to the neighborhood of

this is

*Not clear; meant to

the range as described

refer to

and mapped does reach the middle Ob' unless 1920-30 period (see Fig. 211) Sci. Ed.
;

v^

03

500
^ V

840

563

Fig. 211.

western Siberia (V.G. Heptner):

cent.;

Borders of the distribution of pine marten, Martes {Martes) martes L. in 1 Border at the beginning of the 50's of the XX Proposed limits of the range Same in the 20' s and 30' s of the XX cent.;

in the

XVII

cent.;

Transgression

south of Petropavlovsk.

of the Vasyugan. Thence, the border of the range passes to the 564 Irtysh at the mouth of the and is directed west, to the north of Lake

Saltaim a

little

north of 56 N.

lat.

Farther on, the border

coincides with the [previous] description (for the distribution of

marten

in Siberia, see

below).

841

There

is

poorly defined information concerning the appearance

of (transient) pine martens far to the south in riparian forest tracts,

along the Ishima around Yavlenka about 100 km above Petropavlovsk (Leninsk region of North Kazakhstan district;
Sludskii, 1953).

The southern border of the range in the European part of the Union in the west covers the forest regions of central Moldavia on the Kodora (Kuznetsov, 1952). Crossing the Dnestr, it includes the
Baity region (former Baltskii
est (west of

.;

Ornev, 1931) and Savransk for-

Pervomaisk), passing toward Dnepropetrovsk (Samarsk

farther, to

pine forest, the Samara river which flows into the Dnepr from the
to the mouth of Ognev, 1931). From the mouth of the Oskol, the border extends to the Don, leaving it around Pavlovsk (Shipov forest) and thereafter, to the Khoper around Novokhopersk (Tellermanovsk grove; Ognev and Vorob'ev, 1923). Along the Khoper, the southern border of the range ascends northward, approximately to 52 (Turka), and thence goes eastward, reaching the Volga at Saratov or a little below (it occupies the forest-steppe on the left bank; El'pat'evskii, Larina and
left;

Yurgenson, 1932) and

Izyum and

the Oskol in Donets (former Izyumskii Co.;

Golikova, 1950).

segment of the southern border of the range, the marten extends into the forest-steppe zone, and its actual outlines are very complicated. The described line passes through
In the above-described

the extreme southern localities, in part separate islands of forest

massifs or along the southern extremities of the riparian steppe

forests. Therefore, its

[delineation] here

is

strongly schematized

and basically of a conditional character. In nature, the border line bends more or

less

northward

in a

Such is the picture between the Dnestr and Bug, the Bug and Dnepr, and between the Dnepr and Donets. The range border withdraws [to the north] particularly strongly in the areas between the Donets, the sources of the Oka, and the Don. Here, from the Izyum region, the border ascends sharply to the north, along the Oskol. Leaving the forestless regions of Maloyaroslavts, Liven and El'ets beyond the range, the border extends to Orel and then turns to the east, through Novosilya region (Ognev, 1931), and crosses the sources of the Don, and goes to Ranenburg (Tupov, 1925) or a little south, and then descends, as an extension, along the Tsna to Tambov.
places, over the unforested steppes of interfluves.

number of

842

From Tambov
it

descends

to

or from the sources of the Don, Voronezh and then along the Don

in

an unclear way,

to Pavlovsk, reach-

ing the previous line.

For all of that, marten distribution in the basin of the Don is extremely sporadic and is associated with the separate forest masses along the rivers or on watersheds. Thus, in the Voronezh district, marten was known from Voronezh preserve (Grafsk forestry allotment) and neighboring forest massifs,
forest (N. Severtsov, 1855;
in

the forests along the

Bityug, and particularly around Bobrov and in the Khrenovsk

V.G. Heptner), around Novyi Kurlak Kamennaya Steppe (Dokuchaevsk Experimental station around Talovaya station), and in the Shipov forest near Pavlovsk (Ognev and Vorob'ev, 1923).
and
in several other places, in the

Beyond
the

the

Volga the border, apparently

starting near the city

of Engel's, goes northward along the floodplain of the Volga to

mouth of
to the

the Bolshoi Irgiz.

From

there,

forests of this river eastward nearly to

it passes along flooded Pugachev, then returns west-

wards

Volga and extends along

this river.

its

right

bank

to the

mouth

of the Malyi Irgiz, rising somewhat to the east in the extreme

lower reaches of
itself

Thence, the border line again directed

northwards, along the Volga floodplain, reaching the bend of

river (N.I. Larina) towards the Ural

the Samara; apparently, the southern border passes eastward along

the

Samara

and joins the aboveit

described border, which passes from Siberia to the Ural [river].

Along the bottom land of

descends a
565
little

the Ural and the rivers flowing into

(Burli and others), the marten goes

below

it

down the (transients known

river to Ural'sk

and

to Gur'ev; Sludskii,

1953). Extending eastward along the Ural river, the line joins the

above-outlined southern border of the Siberian part of the range.

The western border of

throughout
is its

the European-Siberian part of the range

to the Black Sea formed by the state frontier. The Caucasian portion of the marten's range occupies the forest regions of the Main Caucasus range northwards, including the

whole extent from the Barents Sea

foothill forests.

The range apparently

also includes the forests of

the Terek valley lying on the plain. Information on the marten in

the

forest tracts of the

Parabochevsk forest near Shelkovsk station and in the mixed Terek valley (Heptner and Formozov, 1941)
range includes the
(left

relate, evidently, to this species. In the west, the

basin of the Pshekha river

branch of the Belaya), and perhaps

843 extend somewhat farther.

rior forests of

To

the east, the range reaches the inte-

Dagestan

in the basin of the

Avarsk Koisu (Dinnik,

is

1911). Marten also occurs, apparently, in the foothill forests south

of Makhachkala, but positive information on this

absent. Its

absence

is

sometimes even emphasized

there, for

example, in the

Deshlagara region (Dinnik, 1911). The range also includes all of the wooded southern slopes and the foothills of the Great Caucasus eastwards almost to Zakatal and Nukha.
In the Little Caucasus, the range occupies only the western
part of the territory,

namely the northern slopes of the Adzharthe east of Akstafa-Chai and Debeda-Chai,

Imeretinsk and Trialetsk ranges from Borzhomi and Bakurian' to

the Belyi Klyuch.
i.e.

To

the meridian centering on

rare.

extremely

Lake Sevan, this marten is absent or They do not occur in the forests of Karabakh. To

the south, along the Pri-Black sea region (Adzhariya) the range

extends to the state frontier and along the Tavr, exits into Asia

Minor. In

all

remaining part of the Caucasus, the pine marten

is

absent (Dinnik, 1910, 1915; Satunin, 1915; Vereshchagin, 1947,

1959). Old information on occurrences farther to the east, particularly at

Talysh (Radde, 1886; Satunin, 1896) were denied by

latest data

all

subsequent authors. However, the

on occurrence of

this

species in Iran (see below) do not exclude the possibility of their

appearance

in Talysh.

Indications of occurrence in

some

parts of the Tien

Shan

(Shnitnikov, 1925, 1936; Shostak, 1927; N. Severtsov, 1873;

Zarudnyi,

1915; Laptev,

1929) and even in Trans-Baikaliya

(Cherkasov, 1867) are completely wanting in foundation.

The marten

represents a stenotopic forest animal, connected to

a significant degree, with forests of tall trees. In connection with

this, its distribution

maintains a more or less continuous character

only in the northern parts of the forest zone. The farther to the

more it becomes sporadic, in accordance with the relaand dispersion of forest massifs. In the forest-steppe and steppe zones, as has been indicated, its distribution assumes an extreme form of separate islands located far from each other. At the present time, in connection with rapid changes in natural conditions its distribution is becoming all the more sporadic in the
south, the
tive area

north.

In

most parts of the marten's range,

if

we do

not take into

consideration the several reductions associated with landscape

844
changes, as for example, the felling of forests,
not

it

was

not,

and
is

is

now

subjected to evident changes. The described range

all

the

present species range, in

events in the European part of the

Union and

the Urals.

As

for the Trans-Ural

is,

and western Siberia, the history of the

marten's range

apparently,

more complicated.
that earlier the

In the literature,

few assertions distributed beyond the Urals and

there are quite a

marten was not

that the range of the species then

increased eastward (Heptner, 1936, from data of L.G. Kaplanov;

Skalon and Raevskii, 1940; Sludskii, 1953 and others). Concerning this, it is considered that this process was not far back in
time

the

last

century and particularly at the beginning of the

present one, and that earlier, the marten was not present beyond
the Urals.

The reason

for this
in the

be the sharp reduction

566 the

phenomenon is usually considered to number of sables and its exterminathis,

tion in several districts.

With

a suitable ecological niche for

marten was freed-up. With a normal number of sable, the marten

its

could not penetrate into the east because of

conditions

"resistance".
in

It

has

been considered that the cause might be change

natural

a shift in the black taiga through thinning, or both

conditions together.

Analysis of literature records of the 18th cent., half-forgotten

or considered unreliable, and of

new

archival materials (Kirikov,

1958, 1960), shows, however, that in

already

West

Siberia, the marten

it

was

known

a long time ago. Thus, in the 17th cent.,

lived

here along the border of the forest zone and throughout the forest-

steppe in quite considerable numbers and

cent.,

still

remained

in the 18th

though

marten near

counties of that time,

The marten existed not only in Tobolsk and but even in Tomsk, i.e. farther east than now. Moreover, in the northern part of the West Siberian taiga (Berezovsk county), marten was absent in the 17th cent.; it was
also absent in the middle section of the taiga zone

.
in

very small numbers. Pallas also took note of the

in

Surgutsk

county (1630, 1650, 1660) or

was met with here very rarely (Kirikov, 1960). Evidently, marten was distributed in southern Siit it,

beria far to the east, but in the north,

the [Trans-Ural] plain.

apparently, did not reach

Apparently during the 18th and 19th

cant part of

cent.,

marten (as well as

difficult to

sables) completely or almost completely disappeared in a signifiits

previous West Siberian range.

It

is

understand the complete disappearance of information about this

845
species in the literature of the 19th cent., or, on the contrary, the
reference to
its

appearance (settlement)

in

western parts of the country

(Pelymka; Slovtsov, 1892 and others) at the

end of the century.

the

On

the basis of

all

these referenced materials, the history of

the marten range in western Siberia

more correctly presents

following aspect: in southern regions

along the southern part of

from time immemorial (fossils are known 2,500-8,000 years in age; Laptev, 1958) and reached Tomsk district. In the north of western Siberia, the marten was absent. Then, in connection with its destruction in subsequent centuries, the range of the marten contracted very
the forest zone, the marten lived in the forest-steppe strongly,
it

disappeared or almost disappeared in western Siberia.

is

In recent times, the range

being re-established, especially intenfull

sively in the last decade.

still

However, the
it

range in the south

is

is

not re-established. In northern Siberia, the marten

regions, where, previously,

settling

in

new

was

absent. There, coloniza-

tion has been taking place, in our day, especially intensively in the

30' s. Therefore, externally similar

differences.

phenomena have fundamental

detail

for these changes in range are not understood in and require careful investigation; evidently, both of the abovementioned phenomena biocoenotic (relationship between sable and

The reasons

marten) and changes of the landscape are significantly involved. In

the south, where the marten lived earlier, the extermination of

sable and decrease of

its

cant; in the north, equally with the biocoenotic factor,

the character of forests

numbers were, probably, more significhange in was, apparently, the main factor. The re-

placement of sable by marten under certain conditions in some regions is definitely known (Cherdynskii Ural and other places). Naturally, all these events went forward in a background of exploitation, its greater or lesser intensity,

and changes

in its form.

Geographic Range outside the Soviet Union

The range extends through Europe;

in the west,

it

includes Ireland

and England; in the north the Scandinavian peninsula to tree line; and in the south northern spain (Pyrenees), the Balearic Islands, Sardinia and Italy, Sicily and southwards in the Balkans at least to northern Greece.^'* In Asia, the range occupies Asia Minor

'^In Van den Brink (1958) the Balkans are excluded from the range. Marten, however, occupies all of Yugoslavia in the south including Macedonia (Dulich and

Tortich, 1960). There are no data on

its

penetration farther to the south.

846

and northern
system,
it

Iraq,

and to Iran

in

two extensions

along the Elburz

China, where

extends to the meridian of Budzhnurd [Bujnurd] in Khorassan

and from Kurdistan and Luristan southeastward along the Zagros mountains approximately to the meridian of Shiraz (Misonne, 1959).
Information about
the pine marten
is
is
its

occurrence

in northeastern

supposedly even more

common

than sable, and

sold as a low quality sable in the former Manchuria and China

567

Fig. 212. Species range of pine marten,

Martes (Martes) martes L. V.G. Heptner.

847
567

(Sowerby, 1923)

is

deprived of any foundation. The reference

Lukashkin and Zhernakov, 1934). References to the occurrence of marten "in Siberia, Turkestan to Manchuria" are also improbable (Brass, 1926)
applies, of course, to the true marten (see in particular

(V.H.).

Geographic Variation
Geographic variations of pine marten inhabiting our country have been discussed by several authors (Satunin, 1914; Ognev, 1931; Kuznetsov, 1941; Yurgenson, 1947). These studies dealt with both the entire range as well as its separate parts. The scheme, established about 20 years ago, has not been modified since then, and
cannot be considered completely satisfactory. Some populations, characterized by highly subtle characters were given separate names.

These features do not reflect equal representation of "good" subspecies, and several are a priori doubtful. The characteristics of marten skins as a commodity fur, apparently, served as the stimulus for distinguishing

some forms.
is

A
essary.

new

revision of geographic variation in the species

nec-

568

The scheme of B.A. Kuznetsov (1941, 1952) is presented below with a few modifications, critical remarks and nomenclatorial changes. The diagnosis is given in brief format, sometimes with supplements from other authors. 1. Western European pine marten, M. (M.) m. martes Linnaeus,
1758 (syn. sylvatica,
Size large,
tail

sylvestris, abietum).

long. Skull large and massive.

Fur on spine beautiful dark-chestnut color; guard hairs darkbrown, underfur bluish-gray, tips lighter and paler. On sides, fur slightly lighter. Withers and upper part of neck and head somewhat darker than spine. Muzzle brownish, slightly lighter posteriorly. Throat patch usually bright yellow. Belly dark brownish,

darker than sides of body. Base of

its tip.

tail

same color

as back, to

brownish toward

Feet dark-brown. Pelage fluffy and fine.

length of males

males 400-450

480-530 and to 550 mm, of fe250-280 mm, females 230-260 mm (Schmidt, 1943; Middle Europe). Condylobasal length of male skull (22) 81.0-M 86.26 0.4688.0 mm, of females (9) is 77.0-M 79.00 0.70-83.0 mm; zygomatic width of males 46.0-M 48.82 0.67-53.8 mm, of females 42.3,

Body

length of adult males

mm;

tail

848
45.01 0.70-49.2 mm; skull height of males is 30.8-M 32.77 0.25-33.7 mm. Weight of 100 skins 10.5 kg (Kola Peninsula). In western European part of the USSR eastward to Kiev,
Bezhitsa, Smolensk, Vitebsk, Leningrad; Kola Peninsula and northern Kareliya.

Outside the
region.

USSR western

Europe, except the Mediterranean

The
ern ones

identity of
is

Kola martens with our western and southwest-

highly doubtful.

Kola martens are the most valuable form of our martens. 2. Middle Russian pine marten, M. (M.) m. ruthena Ognev, 1926. Dimensions smaller than in preceding form. Skull smaller and
less massive.

Prevailing color light reddish-tawny ("reddish sandy" in termi-

nology of

furriers).

Back

also such color, guard hair reddish-tawny,

tips.

underfur light bluish-gray with reddish-tawny hair

back, but with pale hair

Sides lighter

and grayer than back, underfur on the sides also such color as
colored like back;
tips. Belly darker than sides, tail at base end reddish-brown. Distal part of legs darktawny. Throat patch reddish. A small number of marten are found with chestnut guard hairs and light-blue underfur. Body length of [adult] males 415-M 433-452 (and to 500

at

tail 195-M 218-260 (Yurgenson, 1947). Condylobasal length of male skull (32) 75.3-M 80.19 0.4284.8 mm, of females (18) 71.1-M 74.71 0.55-77.3 mm; zygomatic width of males 41.4-M 44.36 0.36-49.0 mm, females 38.2-

mm), length of

mm

42.22 0.32-44.3

mm;

skull height of

males 27.4-M 31.07

0.14-33.3
In

mm,

females 27.0-M 28.84 0.28-31.1

mm.

middle districts of the European part of the USSR south to the range border (steppe) and on the north to the Vologda-Gor'kii-

Kazan
3.

line.

Absent outside the USSR.

Northern pine marten, M. (M.) m. sabaneevi Jurgenson, 1947
(syn. borealis).

Dimensions, and characteristics of skull as in preceding form. Fur on back in most cases light grayish-tawny, guard hairs
light-tawny without reddish
tint,

underfur

light,

ash-gray with pale

tip.

hair tips. Tail tawny, lighter at basal part and darker at

Legs

dark-tawny. Throat patch light-creamy or pale-yellow, rarely pure

white. Pelage fluffy and soft.

849
569

Dimensions similar
of upper Pechora (67)

to preceding form.

Body

length of martens

380-M 433-460 mm; tail length 170-M 198-235 mm. Weight (42) in winter 580-M 784-1090 gm
In

(Yurgenson, 1947). Weight of 100 skins 8.5 kg.

European
line,

part of the

USSR

north of the Vologda-Gor'kii-

Kazan

except northern Karelia and Kola Peninsula, northern

half of Urals (north of Sverdlovsk meridian), and

West

Siberia.

Absent outside the USSR.

Systematic relationships of this and the Middle Russian form
are not completely clear (distribution after B.A. Kuznetsov).
4.

South Ural pine marten, M. (M.) m. uralensis Kuznetsov,

1941.

Dimensions of body and skull large, near those of western European marten and larger than those of Middle-Russian. Predominant color light grayish-tawny tone with highlights of reddish tones on rear part of back. Guard hairs on back light grayishtawny, underfur light-gray with whitish or pale hair
tips.

Withers

somewhat darker than back, sides lighter than back with whitish underfur showing through strongly. Throat patch very light, almost white to often white. Ends of feet brownish, tail base same color
its tip dark-tawny. Fur fluffy and soft. Condylobasal length of male skull (7) 79.8-M 83.57 0.7887.0 mm, female (2) is 74.9-75.0 mm; zygomatic width of male 45.0-M 47.14 0.43-49.1 mm, female 43.0-M 43.6-44.2 mm; skull height of male 31.8-M 33.0 0.41-34.9 mm, female 29.4-

as back,

30.5-31.6

mm. Weight

of 100 skins 9.5 kg.

In southern Trans-Volga, Bashkiria

and Urals south of the

Sverdlovsk meridian (B.A. Kuznetsov).

Absent outside the USSR.

Systematic relationships of the Ural marten with the Middle

Russian and northern ones

are, in

some

respects, not quite clear.

The South Ural forms

are characterized

posterior part of the back which are not developed in the other

by reddish tone on the two

forms (B.A. Kuznetsov).

Caucasian pine marten, M. (M.) m. lorenzi Ognev, 1926. Dimensions very large, skull large and massive. Color dark-tawny with reddish-olive tint. Guard hairs on back
5.

tawny, undercoat bluish-gray, moderately dark, hair

yellow. Sides lighter than back with a grayish
tint.

tips

sandy-

Belly brownish-

tawny with reddish

strip

along the mid-line. Feet dark-brown. Throat

850
light-orange or bright-orange, rarely yellow. Basal part of
chestnut, tip brownish.
tail

dark

Body

length 50-58 cm,

tail

length 23-25 cm, and [hind] foot

length 8-9 cm.

Condylobasal length of male skull (38) is 43.1-M 82.36 0.62 (85.3 mm)35-53.0,* females (14) is 74.6-M 75.43 0.45 47.42 0.45 (78.3 mm)-81.2; zygomatic width of males 43.1,

mm

mm

mm
mm;

(50.9 mm)-53.0, females

skull height of

41.2 M 44.14 males 28.4 M 31.87

0.50 (43.7)-48.3 0.33 (32.1

mm)

-34.0
In

mm,

females 28.1-M 29.93 0.31 (29.5)-31.8

mm. Weight

of 100 skins 11 kg.

Middle Caucasus and Trans-Caucasus.

Outside the

USSR in

contiguous parts of Turkey and Iran.

well characterized race.

Outside the boundaries of our country, the following forms are

usually recognized:
1)

M. (M.) m. latinorum Barrett-Hamilton,

1904

Italy (except southern), Sardinia, Balearic Islands; 2)

M.

(M.) m. notialis Cavazza, 1912

570

South

Italy south

of Abruzzi (V.H.).

Biology
Population. The pine marten, restricted in
forests of the
its is

distribution to the
relatively

European part of the USSR,

few

in

numbers. The entire size of the population of this species before the October Revolution was approximately 40 thousand individuals or a little

after the

more. The number of pine marten greatly increased October Revolution. Very approximately, its numbers may
equal to 200-300 thousand.

now be considered

Some

estimations

of numbers of the pine marten refer to tannery figures for the

periods of irregular exploitation (1922-1925).
ritory

The Northern Terat

produced about

21%

of the annual catch of marten skins;

that time, the western regions

were considerably richer in martens than the eastern. The Urals produced about 16-17%; montane regions of the northern Caucasus 11%; Leningrad district 4%. These four

harvest regions produced about

52%

of the entire catch of the

USSR.

''Average dimension of adult marten (males and females) in West Caucasus

(from Ryabov, 1958). Maximum and minimum values, male condylobasal length, clearly
small.

in error;

too

may be

correct

Sci. Ed.


851

The

forest marten

is,

within

its

range boundaries irregularly

suitability of habitat

distributed, a fact conditioned

in the different regions

by the percentage of forested areas

and refuge

and the degree of

(nutritional conditions, availability of prey items, nest

conditions) within the forest tracts.

Of no

small importance are the

relationships (relative weights) of lands of various values which

are inhabited

by the species with various

all

densities.

Habitat. In

features of

its

structure and ecology, the pine

it

marten

is

closely associated with forest. Within such forests,

reveals very significant plasticity in relation to external environIt is encountered from the sparse pine forests of Kola Peninsula and northern taiga to the oak forests of Mediterranean type, the "maquis" of Sardinia and tall beech forests of

mental conditions.
the

Adzhariya and Kolkhida. Within the limits of such landscape zones are the most preferred habitats of the pine marten. In the coniferous forest zone,
is
it

obviously attracted to dark coniferous forest. Most often these

are old, large-trunked, multi-layered spruce stands in hilly regions.

Less preferred, because food-poor and less suitable for nesting, are
"even-age" spruce, with their single-age, most often slender, trunks. In the pine forests, the number of martens is not distinguished by stability because of instability of food reserves and the ease with

which marten are destroyed by hunters. In 1952-1958 in Arkhangelsk district, in the spruce forests the tracks of pine marten were met with twice as often as in pine forests and other
plantations.

series of daily trail surveys

showed

that

the total length of trails are found in the spruce forests,

in pine forests

81.5% of 11.4%

and only 7.1%

in

in other plantations.

butions

changed

the

different

seasons

These distridepending on

characteristics of distribution of the food resources.

In the zone of

mixed

forests, the pine

marten prefers spruce-

broad-leaved forests, oak groves, linden forests, and

spruce forests

among

the

complex

spruce forests, the spruce-aspen forests

and the so-called pine-spruce aggregations (pines, spruce-mountain sorrel, and spruce-bilberry). In the zone of the broad-leaved forests, it prefers the large broad-leaved forests and their combination with

montane pine
limits.

forests, in the

montane

forests of the north-

ern Caucasus, the fir-beech and beech forests, and the forests lying
at their

upper

Class

quality areas here are the

fir

forests

of the lower, middle and sub-alpine belts (usually these are mixed

852
fir-beech or beech-fir forests). Class
pear, chestnut, aspen
II

quality areas are oaks, beech,

in the

and alder groves

broad-leaved [forest]

zone (Ryabov, 1958).

Quality (bonitet) of habitat for pine marten regularly increases

571

from north to south parallel to the change in forest biogeocoenoses and simultaneously with shifts in the nutritional regime. The ecological optimum of the species lies in the montane forests of the northwestern part of the Main Caucasus range. This demonstrated regularity is determined, first of all, by the feeding habitat. Food. The pine marten belongs among the group of polyphagous carnivores, and therefore its existence does not depend upon the abundance of any one type of food. The following are among the most important foods of pine martens:^^ 1) mouse-like rodents
(mainly red-backed voles); 2) squirrels; 3) birds; 4) insects; 5)
forest fruits (including berries

and

nuts).

Fig. 213. Pine marten.

Caucasian Preserve. Photograph by L.S. Ryabov.

"Occurrence

in

food of

20%

or more.

853

and species compostition of the food is highly variable. Each locality and each season is characterized by its own food set and by the relative weight of each. From year to year they also do not cease to vary. Nutrition variation depends on

The

relative weight

which all the more determines the specific composition of the food and on the time of year, which are connected with the presence of food and the degree of its availability for pine marten, and finally on the abundance (yield)
the geographic situation of the locality

of each food type.

carnivory of the increases from south also reflected south. This and omnivory from north
marten foods
ation of the
diet
to
is

There

is

a definite regularity in geographic variation of pine

to north,
in vari-

chewing apparatus (Yurgenson, 1951). From north to and birds, especially tetraonids, as well as bird mammals south,
in the

eggs regularly decrease

food of the pine marten.

On

the

contrary, occurrences of mouse-like rodents among mammals, and of insects increase from north to south. In the coniferous taiga, plant food is of more frequent occur-

rence in the snowless period of the year.

On

the contrary, in the

zones lying more to the south

yearly occurrence
is

is

in the

snowy

period.

Its

average

similar everywhere and reflects the plant food

requirement of the animal.

572

Seasonal variation of food

riod,

also well illustrated. For

example
pe-

the squirrel mainly occurs in pine marten food in the

snowy

constituting about

snowless period

occurrence or more while in the not more than 6-8%. The same may be said

44%

about tetraonid birds and mountain ash berries. On the contrary, insects, bilberry and mouse-like rodents are predominately typical food in the snowless period, while in the snowy period, they, except
the latter, disappear, or are

There
size

is

marten. In the

met with significantly more rarely. marked sexual dimorphism in foods of the pine food of the weaker females, which are of smaller
i.e.

and weight, hares, capercaillie, black grouse

the larger

prey are not met with.

On

the contrary, hazelgrouse and

mouse-

like rodents are more frequently encountered in the female's food

than in the male's (Yurgenson, 1947; Gribova, 1958).

Data of geographic and seasonal variations

of pine martens were used:
1)

in

foods are pre1948);

sented in Tables 57 and 58. In these, 2,751 data points on nutrition

Lapland

587 (Nasimovich,

854

upper Pechora 593 (E.N. Teplova and P.B. Yurgenson)*; Vologodsk district 205 (Gribova, 1958); 4) Zhiguli 151 (P.B. Yurgenson)*; 5) the middle zone 220 (Grigor'ev and Teplov, 1300 feces and 1939; Yurgenson, 1939); 6) Northern Caucasus 54 stomachs (Donaurov and Teplov, 1938; Ryabov, 1958). From year to year, depending upon the abundance and degree of availability of one or another food, its occurrence in the food fundamentally changes. In the upper Pechora (Teplov, 1960) for
2) 3)

10 years (1937/38-1948/49), the occurrence of basic food types

574

changed within the following limits (Table 58). In Table 59 only the average occurrence of the different components of the food by seasons is given.
Table 57. Foods of pine marten of different geographical regions for the whole year (% of occurrence)

Food type

Lapland

Upper
Pechora

Middle zone
48.7
24.7
11.6

Zhiguli

Northern Caucasus
62.3 58.3

Mammals
Mouse-like rodents
Squirrel
Insectivores

70.3
34.3
3.1

63.5

83.0

34.9
26.3
6.7

72.0

4.4

4.9

Birds

34.8

30.6
15.7

21.5
7.4

Including
tetraonids

23.0
6.9
10.5

Bird eggs
Insects

5.7

4.1

4.0
24.0
14.6 12.8

9.3

3.0

18.0

0.6

10.7

7.4

39.5
14.8 14.8

Plant food
Fruits (including

23.9
17.5

25.5
11.6

No No

data data

berries)

Table 58. Foods of pine martens in the upper Pechora over 10 years

Food type

of occurrence

Average,

M inimum
Voles
Squirrel

Maximum
29.4
58.1

7.8
13.7

23.3

36.8
15.1

Tetraonid birds

4.0
0.0
5.0
6.1

24.0 23.0
39.1

Other birds
Plant food

8.3

24.5
11.8

Other food

20.9

"No year given

Sci. Ed.

855

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oo oo Tt (N Tt 0\

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r^

r<^ vri

Tj-

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5 Z

U ( 3 2

CN cn -O f^

1 ^
I

Tj-

>n oo in in
I

--

>n >n

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fS

Z Z
^
i;

'

m ^

vn 00

>

^
00 in so vb 't

>
r}-00(N

Tt >OON400r~-

r-1

OO On r-^ SO

9
^~
.;

=>

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i=

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XI aCO 00

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9 u 5 ^ 3 ^

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856
In Arkhangel'sk oblast,

the occurrence of different foods

from 1949-1961, i.e. during 13 years, was as follows (Grakov, 1962)

(Table 60).
In

one of the provinces of Sweden (Heglund, I960)*, the charas follows

its variation (%) within a three-year pe(1956/57-1957/58-1958/59)

acter of winter food and

riod

were

Mouse-like rodents
Squirrel

44.9
39.7
7.7

(25.0-61.5%) (25.6-60.7%)
(6.5-9.0%)

Passerine birds
Birds (general)

21.8
7.7

Bird eggs
Fruits (including berries)

12.8

The occurrence of

squirrel in

marten food increased with the

is

decrease in mouse-like rodents and vice versa.

From

the tables presented,

it

evident that even within the

boundaries of one zone (in the European northern taiga), considerable variation in the degree of occurrence of the main food groups
is

observed. In particular, the range of variation in the separate

is

food groups
in

more sharply marked

is

in

Arkhangel'sk

district than

Pechora, because there (in Arkhangel'sk district) the occurrence

lower.

of squirrel and plant food

The

latter results

from the

absence of nut pine "nutlets"

sula the differences are even

in winter food.

On

the Kola Penin-

more marked.

Table 60. Nutrition of the pine marten of Arkhangel'sk oblast during a period of
13 years

Food type

Occurrence,

%
Maximum
75.5
17.5

Average (m)

Minimum
Voles
Squirrel
12.5

45.1
7.1

White hare
Insectivores

9.2
6.1

5.0
2.0

Birds
Including tetraonids
2.8

32.5 22.8
10.5

21.5
13.6
3.4
3.5

Bird eggs

Frogs
Berries

9.2
43.1

11.2
3.2

Mountain ash

17.0

*Not

in Lit.

Cite--Sci. Ed.


857

Mouse-like rodents occur as the main food of pine marten

everywhere, mainly voles, and among the latter, the bank [Clethrionomy glareolus] and red-backed [C rutilus] voles. Only in the northwestern Caucasus, are they replaced by local species bush voles [Pitymys spp.] and others. In the food of pine marten, mouse-like rodents are found even when their number decreases in the forest. High numbers [of voles] in the forest causes the concenmarten in biotopes that are less characteristic for them, such as in glades, burns, along forest borders, etc. This situation also causes intensification of the autumn migration of
tration of pine

young martens. Mouse-like rodents, especially together with

secondary and occasional foods, ensures minimum nutrition for martens in famine years. In the snowless period of the year, when
the capture of mouse-like rodents
tens, their relative
is

considerably easier for mar-

weight

in the

marten's food increases.

their species

Species composition of mouse-like rodents utilized by martens

is

variable.

It

depends not only on

composition and

relative

abundance

in the local forest fauna, but also

on

their rela-

575

more active wood mouse [Apodemus] is always less common. The forests of the Black Sea coast of the Caucasus constitute an exception where, in some places, other
tive availability: the

mouse-like rodents are absent.

widespread belief that the pine marten feeds mainly on squirrels. From the tables presented above show that this is not the case; the pine marten can exist very well, where squirrels are completely absent, as for example in the Caucasus,
In the past, there

was

where the marten

thrives.

Even now,

after the establishment of

squirrels in the forests of the northwestern Caucasus, the pine marten

makes

little

use of this resource (6.6%). Moreover, in the taiga

forest during the snowless period

when food

is

significantly

more

available for marten than in winter, the importance of squirrels in

its

the nest

food sharply decreases. The destruction of young squirrels in by martens is still not based on facts. In the last years, the role of the pine marten (and also of the

sable in Siberia) in reducing squirrel

numbers has been strongly

elucidated (Yurgenson, 1954; Ivanter, 1961; Grakov, 1962; Yazan, 1962). The relationship between the number and density of both
species per unit of forest area, as well as the degree of abundance

and availability of the marten's main food are of fundamental importance. On average, the Pechora marten (which feeds more on

858
squirrels than

anywhere

else) eats

8-10

squirrels per winter. In

are,

few and martens may destroy up carnivores these abundant, hand,

years

when

squirrels are very

on the other
of
all
is

to

30-35%

squirrels; in other years, the importance of squirrels

consider-

ably lower.

It

is

believed that in the European North (Grakov,

squirrel
in local

1962) the pine marten lacks the power to affect the dynamics of numbers and cannot determine their number. Marked losses
populations of squirrels are only caused by marten in those

rare years

when the low number of squirrels is accompanied by a of the tetraonid birds and mouse-like rodents, while number low martens are, at the same time, numerous.

An abundance
in their

of squirrels in nature does not cause an increase

in

voles.
in

marten food, especially if there are many Pechora, increased occurrence of squirrel marten food occurs in connection with the increased occurrence
occurrence

As was noted
of

in

in nature

weak and

sick squirrels.

marten of the European taiga and some parts of the mixed forest zone, tetraonid birds acquire an essentail importance, especially capercaillie [Tetrao], black grouse
In the nutrition of the pine
tetrix], hazel grouse [Tetrastes bonasia] and willow grouse [Lagopus lagopus]. Among these birds, the hazel grouse is the most frequent prey of marten. Tetraonid birds are hunted by marten mainly in their snow burrows and night resting places and, therefore in summer, their occurrence in marten food decreases

[Lyrurus

significantly and

becomes more or
it

less accidental.

By means

of

tracks in the snow,

ful

is

frequently possible to trace the unsuccess-

hunts by pine martens of capercaillie or hazel hen. The quantity of tetraonid birds in marten food usually increases in the years with a reduction in numbers of voles and squirrels. In Pechora,
capercaillie and black grouse are only found in the food of the larger and stronger males. In the food of females, only hazel grouse

are observed (Yurgenson, 1947). This also occurs locally in the

Vologodsk

district (Gribora, 1958).

the food of males. In

White hare also occurs only in the Central Forest Preserve, there were indi-

vidual male-hare hunters specializing in tracking and catching hares.

Their daily

trails

were often particularly long.

birds
is

The capture of small

a characteristic feature of pine

marten although their relative weight in the ration is quite small. This is determined by the small size of the prey. Among the birds
hunted are hole-nesters: woodpeckers,
tits

and nuthatches.

859

Shrews

in

marten food are either an accidental element or an

indication of sharp deficiency in principal food. Sometimes,

com-

mon moles

[Talpa europa] appear

in the

food of martens
is

in certain

winters and in fairly great numbers. This

connected with the

death of moles through strong freezing

of the taiga soil in years

tetraonid birds, mainly

with

little

snow (Teplova,

1947).

The eggs of

those of capercaillie, in winter food of marten

576 cial conditions. This is

is also due to speconnected with the death of the embryo in

the shell early in spring, due to the excessive cold, a situation

which occurs only when the female leaves the eggs for a long time in the years characterized by the repeated occurrence of freezing weather throughout June (Teplov, 1948, 1960; Semenov-TyanShanskii, 1959).

Sometimes, remains of frogs and

winters with food deficit,

their

eggs are found among

the remains of marten food in winter. This is observed in those

when

the carnivores find,

somewhere

in

a thawing rivulet, an overwintering accumulation of frogs. In Tatariya, on the Malyi Cheremshan, tracks of martens were fre-

quently observed proceeding along the river

ice,

where American

mink hunted

frogs. Usually, the

mink

left

the frogs on the

snow
snow-

and the martens took them off (V.A. Popov).

In the southern parts of the species range, marten in the
free period eat a great quantity of insects, including wasps, bees,

bumblebees, beetles (especially ground beetles). In the taiga and in middle zone forests, wasps, bumblebees and honey bees are often
encountered. In the Caucasus, beetles predominate. In the taiga,
the significance of this food group
is

naturally less, as the food

objects are fewer.

On

discovering a tree hollow occupied by wild

bees, the pine marten willingly and protractedly feeds on the honey

and the bee larvae.

Plant foods are met with in taiga forests, chiefly in the snowfree period. Then, first place
is

occupied by bilberries. Other

fruits,

including even choke cherry are of secondary importance. Only in

the upper Pechora do nut pine "nutlets" acquire very great
significance in

autumn and

winter, naturally in years with a yield.

They

are usually found in martens' stomachs together with forest

voles. Marten feed on mountain ash berries in winter far beyond correspondence with their presence and availability. It utilizes ash
fruits

quite moderately, especially

when
is

other food

is

found

in

great quantities. There,

when

nut pine

absent in the taiga, ash

860
fruits

and other plant food appear in marten food in winter when few (Grakov, 1962). In the southern parts of the range, fruits, especially in autumn and winter, have greater importance for marten than in the taiga. In the northwestern Cauprincipal foods are

casus, together with a

tain) ash,

whole list of wild fruits (including mounthose of yew, which are poisonous to man, are of essential
is

importance.

Polyphagy
It

a very characteristic feature of the pine marten.

one food, to turn to another. However, the comparison of combinations in one stomach of various numbers of foods with the degree of fullness of the stomach and the fatness of the animal, shows that differences in daily rations is a negative feature, indicating insufficiency in the main full-value foods. The analysis of multiyear data on food of pine
enables
it,

in case of a deficit of

marten also shows that along with the existing relationship between the degree of abundance of basic foods in nature and the frequency of their occurrence in food, another relationship is
present

decrease

in the

occurrence of one food group causes an

increase in

group or groups, independent of the natural abundance of these foods in nature. On the whole, as a result of polyphagy in pine martens, deep and prolonged de-

demand

for another food

pressions in their numbers are not observed, especially in those

places where the conditions are more favorable for their existence. However, though rarely, there are seasons when a majority of the main foods in nature exhibit low numbers and martens starve. From Table 68 on foods of the pine marten it is evident that
in the

snow-free period, occurrence of mouse-like rodents, mainly

forest voles, bird eggs, insects

and

their larvae as well as fruits,

obviously increases in connection with more varied and more available food assortment. At the same time, the occurrence of the more
difficult to obtain foods falls off: squirrels

has been already

577

and tetraonid birds. It from this general Thus, in Arkhangel'sk deviations. local some situation, there are were found to voles food district (Grakov, 1964) in summer constitute 70% of the data, birds 23.2%, insects 24.2%, fruits 21.2%, and among birds, the importance of (including berries)

mentioned above

that apart

tetraonids declines and the role of the small birds grows, lizards
also appear, and the role of shrews
squirrel decreases.

food of

last resort

and

861

The stomach of marten holds

of the live weight of the animal
nature.
It is

a quantity of food equal to 1/10

this is the

optimal daily

norm

in

stomach contained 60gm of more often 90 food, but about 50 gm. The pine marten cannot eat more than one squirrel per day and, quite often, leaves a part of the carcass. For estimating the general provisioning of marten with food in a given year and season, the investigator is led by data on the average fullness of the stomach with food (by weight) and the number of empty stomachs. In the middle and northern taiga zones, marten feed less well than in the mixed forest zone. The average fullness of the [northern] Pechora marten stomach for seven winter seasons was 28.7% of optimal fullness and in the middle zone 80-95%, which constituted 50-70 and up to 90 gm. In Tatariya (Grigor'ev and Teplov, 1939), fullness is about 44% (32 gm). This all accords with geographic variation in marten nutrition in different sections of their habitat and the duration of the daily route. In Pechora marten, the average percentage of stomach fullness in winter for a seven-year period ranged from 14.6 to 51.1%, and the average weight (excluding empty ones) from 10.6 to 37.1 gm. In Vologodsk pine martens (Gribova, 1958), the average weight of stomach content fluctuated in four winter seasons from 25.5 to 35.5 gm (average 29,3 gm); maximum was 126 gm (a frog and its eggs). In the upper Pechora, better fullness usually occurs with a predominance of squirrel and wildfowl, but at this time, the greatest percentage of empty stomachs is also encounnot often achieved.

The

fullest

tered.

Home

range.
its

From
whole

the time of dispersal of the young, the pine

life

marten spends

within the boundaries of a quite clearly

limited section of the forest

its

home

range.

The

territories

of

males and females most often adjoin. The

until

litter

usually remain

autumn within the mother's territory. With plentiful food, litters sometimes do not disperse in winter. The dimensions of the home range directly depend on the quality of area located within its limits, and on the ratio of various types of lands lying within its limits as well as on the food supply in the
given year and season. Territory and

home

range dimensions are

geographically variable; they regularly decrease from north to south.

For martens of the Kola Peninsula, their size ranges from 15-20 and upto 50 km^ and for northern Urals (Uk-yu valley), 6 km^. In the spruce forest massifs of the southwestern part of Kalinin


862
district (Central Forest Preserve),

it

averages 6 km^ (from 3.9 km^

km^) (Konchits, 1937; Yurgenson, 1937, 1939). In Babkovsk forest tract (on the border of Penzensk and Tambovsk districts)
to 9.2

4.4

in the

km^ (Yurgenson, 1939), in Zhiguly (P.B. Yurgenson), 4.0 km^ montane forests of the northwestern Caucasus, the average

area of the daily

home range
fir

(11 cases)

is

1949; Ryabov, 1959), in

forests

from 32

46.7 ha (Donaurov,
to

72 ha. In certain

from 300 ha. The territory of the home range is assimilated by martens in succession, and the same range may itself comprise places periodically visited by the marten in different hours of its diel activity
cases, martens lived in isolated forest islands with an area of

90

to

period, or else, as

its

hunting area. Individual parts of the entire

home range
marten
is

are not used for hunting and only serve for

shelters.

movement.

Burrows and

The most preferred

shelter of the pine

a hollow tree. In the zone of broad-leaf forests, an insuf-

ficiency of these hollow trees does not usually occur. Here, the
forest marten settles in tree holes of different sorts with a diameter

and more. The opening of the inhabited hole mostly m 4-5 m; in the northern Caucasus at a height of 10-20 m and more), but in certain
of 30

cm

occurs high above the ground (from 2-2.5

578

cases, the entrance to the hole

is

situated in the

main trunk;
is

the presence of light-penetrating cracks in the hollow

not an

obstacle to use.
In the zone of mixed forests, hollows inhabited by martens most often occur in long-standing thick-stemmed aspens, more rarely in those fir trees devoid of tops (crowns), or in other types. In the taiga zone, inhabited holes are found in the nut pine, spruce, fir and aspen. There, where tree hollows are rare (mainly in coniferous forests), the pine marten willingly occupies squirrel nests (dreys), more often, as a form of temporary shelter; having caught and eaten a squirrel, the marten lies down in its nest to rest. In rare cases, the female and her litter occupy the drey (Ognev, 1931). In the middle [forest] zone, it was observed that dreys are more often inhabited by small females. This is not observed in the Pechora basin. Squirrel dreys are mainly used by martens in the first half of winter. Thus, in Pechora basin in autumn, and in periods of little snow in winter, the martens use holes and dreys of squirrels. Marten very rarely conceal themselves in logs lying on the ground. In the second half of winter with heavy snow and cold, (from

863
January) the marten leaves these shelters and moves to the logs

hidden
often

in thick

snow where

it is

safer and warmer. In

Arkhangelsk

oblast (Garkov, 1964), also in the first half of winter, they

more

(57-95% of

cases) use squirrel dreys and tree holes and in

the second

they

use wind-falls and spaces between roots (71-

81%). When approaching a shelter, the pine marten rarely seeks to confuse its track (20.5% of cases), but approaches the shelter through the trees for not more than 50 m. In the Caucasus preserve (Ryabov, 1959), out of 16 temporary

were located in hollow fir trees, 2 were in wind-felled and the rest were in hollows of beech, aspens and in windfalls. Nests of litters in this region always occur in tree hollows. In holes with litters bedding, except the remains of rotten wood,
shelters, 11
trees,

does not occur. In the Tatar Republic, the marten sometimes occupies logs placed in the forest for bees especially in second-growth,

where hollows are few (B.S. Popov). Very

establish themselves in rock clefts (Zhiguli).

rarely, pine

marten

In the winter time, small stores of food are

ered in holes. In such a way, holes inhabited by bees

considered unique *'larders" of this

their

sometimes discovmay be animal; the marten feeds on

tracks to such holes.

honey

for a long time,

making beaten

Daily activity and behavior. In contrast to sable, the pine marten

is

an animal with a distinctly demarcated twilight-nocturnal type of

diel activity.

During the day,

it is

active only very rarely and spo-

radically;

somewhat more

often in the period of sexual activity; the

in the

young pine marten, however, can sometimes be seen

time.

day-

Leading a semi-arboreal way of life, the pine marten climbs but not always and not everywhere. Its occurrence and movement in the crowns of trees is closely connected with, and the main form of, feeding on squirrels. It captures other types of food mainly on the ground. In the forests of Arkhangel'sk district, when they
trees,

feed mainly on voles, only

trees (of

1%

of

its

diel

movements were among

this

132 km). In the second half of the winter,

occurs
the

somewhat more often (Grakov, 1964). There, where food of

forest stratum is particularly scarce

(montane forests of the north-

western Caucasus), the pine marten almost exclusively leads a

terrestrial

way of

life,

elevating itself into the trees only in search

of a hiding place or shelter, or to eat mistletoe berries (Ryabov,

1959). In a night,
it

climbs trees not more than 2-4 times. The

864

same is observed in the sparse northern forests (Kola Peninsula), where its terrestrial way of life is determined by the considerable distance between the trunks of trees. The arboreal mode of life of
this

animal

is

often associated with coniferous forests. In regions

579

where sable and marten coexist, the latter are found in trees more often. On the ground surface, and especially under snow, it orients itself and moves about less well than the sable. The character of pine marten movement in winter is determined by the state of snow cover, especially that covering the tree branches (kuxta). The abundance of much clinging snow makes the movement in the crowns difficult except in cases when the kuxta is frozen to the branches, making climbing in the crown
easier for the marten.

The length of
abundance and

diel

movements

(tracks)

is

determined by the

availability of food,

and

in winter, also

by the

state

Fig. 214.

Caucasus pine marten

in natural setting.

Caucasus preserve. Photograph by

L.S. Ryabov.

865
of the

snow

cover. Activity of the marten

is

closely connected with

its feeding. When satiated, the marten stops hunting and lies up at rest. In severe frosts or snowstorms, when the hunting yield does

not compensate the organism's outlay, the pine marten does not go out from its shelter for several days, even if it becomes hungry. At low temperature, the marten moves little within a small area, and
lies

down

during the day, with slightly filled stomach.

area of the

As with

home

range, the length of diel

movements
its

(tracks) are closely connected with the quality of land, with

food supply, and regularly decreases from north to south. In the

580

Kola Peninsula, it ranges from 10 to 25 km (16 km (Nasimovich, 1948); in Priozersk region of district (Grakov, 1960), based on data from 36 paths over five seasons, it equaled 8 km. With insufficient food, length of the diel route in the Severnii Territory reaches 22 km, in foodrich years, it usually reaches 5.3-6.0 km and does not exceed 7 km (Semenov-Tyan-Shanskii, 1961). In the upper Pechora, the route has a length of from 2 to 6 km (4.2 km on average), in the spruce from 0.6 to forests of the southwestern part of Kalinin district 5.5 km (3.1 km on average; Yurgenson and Konchits, 1937), and from 0.6 to 3.2 km (2 km on finally in the Caucasus preserve average; Donaurov, 1948). In the northwestern Caucasus (Ryabov, 1959), diel movement is usually in the form of a broken line and rarely closed; the average length of 11 complete paths here is 2 km; from year to year it changes only slightly. Female marten (and sable) are always less active than males. The average length of route is 2.57 km for males and 1.48 km for females; it ranges from 0.60 to 3.24 km. Seasonal migrations and transgressions. The pine marten is a sedentary animal to a great degree. Even deficiency of food does not always oblige it to leave the range it has inhabited for several successive years. At the same time, local migrations by this animal are observed. In certain cases, it was observed that the pine marten follows migrating squirrels and appears in those forests where the latter are concentrated. Attracted by the abundance of voles in forest glades, burned areas or in fields, marten appear in these habitats, which are not typical for them. Dispersal and migration occur primarily in young marten from litters at the time when they begin to break up at the beginning of winter. Adult individuals rarely take part in this. The greater the density of the marten population, the farther the young are obliged
forests of the

on average) Arkhangel'sk

866
to

move

in

search of suitable, unoccupied areas. In particular,

to habitats temporarily rich in squirrels

young relocate

and voles,

or they gather at carrion. Old marten sometimes migrate from one

place to another, their worn teeth requiring them to search for

more available food. With

the range

the dispersion of young, expansion of

may

proceed.

Reproduction.
sexual maturity
is

Up

to

now, the time when pine martens

attain

not completely clear. According to data from

animal farms, the animals mature not earlier than the third year of
life or, as a rare

exception

the

second year.

Of 52 young female pine martens in the second year of life, pregnancy was observed in 35% (Ryabov, 1958). In a 15-month
old male,
enlarged.
litters.

on July

18,

the testes
all

were found
in a

to

be greatly

About 45% of

females

population bring forth

In the Caucasus, the percentage of barren females in the

third year of life

and older

is

18.7%. For Arkhangel'sk

(Grakov, 1964).

district,

barren females comprise

22%

Estrus and mating take place from the end of June to the be-

ginning of August, frequently from the end of July-beginning of

August. In individual cases, estrus

August.
It

is

delayed until the end of

commences somewhat

later than in sable.

Generally, pregnancy (Manteifel', 1934; Starkov, 1947) lasts

236-237 days; from other data (11

days (Starkov, 1947). In kidas*,
it

254-256 and 258-274 continues 280 days (Portnova,

cases),
is

1941). Pregnancy extends through a long period of inhibited

development
ity,

(latent period).

Its

duration

variable.

As

a rule

placetation, being connected with the resumption of sexual activ-

and which

is

suppressed by September, takes place in March.

is

The

so-called false rut

dated to that time,

i.e.

an increase in
at that

excitement of the sexually mature animals which

time be-

come more

active than usual, and the males follow the tracks of

the females. In certain cases,

some deviations

are noticed: thus, in

Pechora preserve, embryos 9 and 31 in length were found in martens killed on 9 and 20 January. This situation, and the presence of sexually immature individuals, explain does not occur
in

mm

why

the false rut

in all martens.

marten impregnated

on 15 July

the

German

Federal Republic gave birth on 5 January after

being transported to Argentina; the duration of pregnancy due to

the latent period
*See pg. 833.

was shortened

to

176 days (Ulrich, 1953).

867

From

placentation to parturition, embryonic development in

all

our species of the marten family takes 27-28 days (Kler, 1941)^^.

On

farms, birth of young occurs 22-26 March (Manteifel', 1934).

in April

(Starkov,

1947), or

In case of early placentation,

parturition should take place in February.

581

The number of young

it

in a litter

ranges from 2 to
is 3.8; in

8,

most often

3-5. Average size (for entire range)

the Pechora basin,

equals 3 (2-4), in
litter

Moscow

district, 5 (3-7). In the

northwestern

Caucasus,

between 2-7, usually 3-4 (Ryabov, 1958). The average size of the brood varies negligibly in different years: for Arkhangel'sk district and Komi ASSR, from 2.54 to 3.0 (according to corpora lutea of ovaries, 3.5-4.0); for Kirov district, it averaged 3.7 for 1959-1960 (Grakov, 1962).
size ranges

Growth, development, and molt. Growth and development of

Weight increment in young marten decreases sharply with opening of eyes (30th to 32nd day), when the animals begin to feed independently. Further weight increases proceed regularly, but from 48 days of age, fethe pine marten take place as in the sable.
in weight: at the age of 10 weeks, 20 weeks, male 700 gm, and female 600 gm; at 30 weeks, male 800 gm, and female 700 gm. At the age of 40 weeks (10 months), male 1050 gm, and female 850 gm. At the age of 15 months, the male attains adult weight. In winter, adult males weigh 1200-1400 gm and in summer, 1400-

males begin to

fall

behind males
at

animals weigh 400 gm,

1650 gm. Females weigh 800-1100 gm and 1000-1350 gm, respectively. Replacement of carnassial teeth takes place from 21 July-8 August. Juvenile fur of the female is replaced by 15 September, and that of the male by the end of that month. Senses of smell, hearing and sight begin to appear in the 7th week of life. At that time also, coordinated movements and the typical gait, jumping, are initiated. Climbing ability develops in the 10th-12th week, when springing capability of jumping develops. Young marten begins to kill subadult rats independently in the sixth month of life, having begun to utilize meat as food at the age of six weeks (Schmidt, 1943;* Herter and Ohm-Ketner, 1954)*. The pine marten molts two times annually in spring and in autumn.

"In badger and otter

about 60 days.

*Not

in Lit.

Cit. Sci. Ed.

868
Enemies, diseases, parasites, mortality, and competition. Dangerous enemies which in some way might significantly affect its numbers, are not known for pine marten. Among the predators

which are sometimes successful in attacks on martens, especially young marten, are wolverine, red fox, golden eagle, white-tailed eagle, goshawk and eagle owl. In Kalinin district, remains of young marten were found in the nest of a buzzard [Buteo buteo].
Neither epizootics nor helminth infections obviously influence

number of pine martens. Only a small number of cases are known of infection with pulmonary helminths (filiaroidosis and kernozomatosis), which caused significant plague among pine martens in the Northern Caucasus and in Georgia, accompanied by a great reduction in their number. Pulmonary filiaroidosis
the

infection and the epizootic caused by

it

were noted

in Severnii

Territory in 1951/52-1955/56
high.

when

the

number of martens was

Of 503

pine martens from Arkhangel'sk district (Grakov,

1964) investigated from 1946-1958, filiaroidosis was found in 55.3% and Skryabingulosis in 48.2%. Maximum intensity of the first infection was noted in 1951-1953, of the second, in 19531954. Seriously infected females die from filiaroidosis more often
than males. Martens infected with filiaroidosis are also found to be
intensively infected with Skryabingulosis. Adult females are in-

fected with Skryabingulosis

yearling of both sexes.

strongly infected.

more heavily than adult males and the Poorly nourished martens are always more

In large series of skeletons and skulls, a series of cases of

healed broken bones of the limbs and caudal vertebrae were found;
in

one case

even a self-healed

bullet

wound

of the cranium.

All predatory animals and. birds inhabiting the forest

considered as competitors of pine martens

[Felis silvestris], sable, stone

red

may be

fox, forest wildcat

marten and others. All birds and animals utilizing mountain ash, bilberry, stone pine "nutlets" and
582 other plant

food of marten

may

also be regarded, to

some degree,

as

its

competitors. Cases of interspecific competition for food which

might have displaced the pine marten from its occupied habitat, or which influenced its number, are unknown. In individual cases, the longevity of marten has reached 14-16 years. Under natural conditions, pine martens rarely attain such an
age and their average longevity
is

much

shorter.

869

The age

ratio in separate populations is determined, first of all,

by the intensity of exploitation. In a series of marten from PechoraIlych preserve (76 specimens. Zoological Museum, Moscow University), marten up to one year old constituted 47%, up to two years old 41%, older than two years 9.2% and those older than 2.7%. In the same preserve, from material of comthree years mercial hunting for 1937/38-1948/49 (309 specimens), yearlings constituted 49.9% on the average (from 44% to 58%). In the following winter, after a winter with abundant food, an increase in the percentage of yearlings was noticed in the population. How-

ever, the rise in the percentage of yearlings

may

indicate not only

the increase in the size of the annual increase but also an increase
in mortality in older ages.

Moreover,
is

in the

time of the harvest, a

taken

larger percentage of yearlings

usually captured than their actual

proportion in the population.

in the

Of 464 martens commercially

upper Pechora (Yazan, 1962), adult individuals of both sexes constituted, for a series of years, 35.4%, in particular, 24.5% males and 10.9% females; juveniles were 64.4%. Among the adult martens, the sex ratio

was

1.0

0.4 and

among

those up to one year

it

was
tuted

1.0

1.7.

On

the whole, males in Pechora population consti-

54% and

females,

46%

(Teplov, 1960).

Thus, considering the prevalence of young matens in the catch,

as a

normal phenomenon, and also

that the

preeminence

in the

harvest of males, the sharp decrease in the percentage of females

in the

population with the transition from yearlings to adults

itself.

(1

year and older) draws attention to

Of 126 martens
[yearlings]

taken in 1939/40 in Volzhsko-Kamsk Territory

in their 3rd year

(Aspisov, 1959), juveniles constituted 51.6%, in their 2nd year

4th year 11.9%, years 2.4%. Individuals above old were absent. In 1940/41, among 289 harvested marten, were year 6.2%, juveniles 70.6%, yearlings 19.4%, quite year 1.4%. fourth year 2.4%, and

30.2%,

in their

3.9%, and

in their 5th

year

five

in their third

in

their

in their fifth

It is

obvious that the percentage of juveniles in 1940/41 increased not

only on account of the higher
fertility,

but also on account of the

decrease in number of older individuals. The average percentage

of juveniles, over several seasons, (594 individuals) was 54.4%,

(42.9-70.6%). Males in this series constituted 61%. Males commercial catch always and everywhere predominated.

in this

870
In

Arkhangel'sk

district

(Grakov, 1964), after winters with

conditions favorable to martens, the percentage of individuals up to one year [juveniles] in the catch increased to 59%, and after

unfavorable winters,

fell

to

43-44%. With favorable conditions,

changed in favor of females (from 56%). In the unfavorable years, the percentage of pregnant females fell as much as 83% (Kirov district, 1959/60). This was
the sex ratio in the population

38%

to

Vologodsk district (Gribova, 1956). With three being the average number of embryos in Pechora pine martens at the beginning of the harvest, for each female there were two juveniles; mortality was 30%*. In the population, the average percentage of juveniles is 48% with annual fluctuations from 35% to
also observed in

58%

(Teplov, 1954).
13 harvest seasons (1949-

In Arkhangel'sk district, during

1961), males exceeded females by 5.8-31%, averaging 7.2%. In 5 years of track counts (49 cases), males were 54%, and females 45%. By means of a thorough, even complete exploitation of one area in the course of three seasons (102 data points), males were

54% and

females

46%

in the catch

(Grakov, 1964).

It

was

estab-

1964) that data from the harvested population sample, because of selectivity during harvest, the percentage of juveniles in the population was overestimated by 14%. Baited traps
lished (Grakov,
583

more often capture males, but hunting with guns assisted by laika dogs, which occurs at the start of winter, takes more females.
In the northwestern

Caucasus (281 martens), juveniles consti-

55.2% (ranging from 49.9-58.2%), yearlings 31.7% (29.138.6%) and 2 year olds and older 13.1% (12.1-13.4%) (Ryabov, 1958). Males were 56.9% (54-58%), females 43.1% (42-45%). The number of males in the first year of life exceeds that of fetuted

males 2-2.5 times. In this way, the pine marten population is quite quickly restored. Population dynamics. The number of pine marten is subjected to oscillations which, however, are relatively small. High levels of populations are observed during considerable time intervals, about
9-1
1

years. Population stability

is

explained by the polyphagy of

the pine marten.

Noticeable increase or decrease in numbers (in the absence of harvesting) is usually determined by the simultaneous abundance

and availability of some of their principal foods, which does not

*Rounded value

Sci. Ed.

871

occur often. This

is

predetermined by food abundance during

The rarity of such combined events determine the rarity of noticeable population changes.
several successive seasons combined.
is compensated for by the abundance of another, or several other secondary foods. Pulmonary infection has a place in depression of numbers, but is rare and does not constitute a universal phenomenon. The most extensive filiaroidosis among pine martens of northern Europe was observed in years of favorable feeding conditions,

Usually, deficiency of one main food

but after unfavorable years. Skryabingulosis, less pathogenic for

pine martens, differed in
its

greater intensity and extent during

years with unfavorable feeding conditions. During the period from

1951/52-1955/56, the extent of filiaroidosis was determined to be between 83.7 and 100%, and Skryabingulosis in the period from 1951/52-1953/54 as 71-87%. A peak population of Arkhangel'sk
pine martens was recorded in the winter of 1951/52, but the great-

number of dead animals was found in 1950/51 and 1954/55. The next rise in numbers was in 1957/58 (Gribov, 1959; Grakov,
est

1962). Helminthic infections are always due to decrease in the organisms resistance as a result of poor nutrition. It is not the ultimate cause of population changes, but one of the links in the process of decline. Harvesting is a very obvious, and sometimes
the prevailing, influence on populations.
ing,

With intensive harvest-

high numbers never occur. Under conditions of absence of

hunting (preserves), the numbers of pine marten fluctuate within

comparatively narrow limits

by

a factor of 1.6 (Pechora; Teplov,

1951), and a factor of 3.2 (southwestern Kalinin district; P.B.

Yurgenson) during 15-20 years.

In Pechoro-Ilych preserve (Teplov,

1960), during the period

from 1938/39-1948/49,
rence of their tracks
in the region of
in

i.e.

for 10 years, the frequency of occur-

pine forests along a 10-km route in winter

i.e.,

fluctuated between an average of 0.5 to 0.95;

by 1.9* fold and

dark coniferous forests, more favorable for the

to 1.3;
i.e.,

pine marten
district

from 0.8

only 1.6 fold. In Arkhangel'sk

this fre-

(Grakov, 1962) during eight years (1952-1959)

to 2.9;
i.e.,

quency fluctuated from 0.7

It is

4.1 fold.

obvious that

stability of population

numbers serves also

as

an indicator of more stable and more optimal habitat conditions

especially in conjunction with a high level of numerical fluctua*

Misprinted as 19

in

Russian original

Sci. Ed.

872
tions. In

Arkhangel'sk
be plentiful.

district, the

pine marten population

is esti-

mated

to

If there

are territories

where the capture of pine marten

is

prohibited (preserves,

many

years of legal protection), the stable

abundance of these carnivores is secured by a mixture of hunting areas, on account of pre-winter settlement by young marten from
dispersing
584
litters.
It

Field characteristics.

is

rare to encounter a pine marten.

One may
logs,

often observe the characteristic feces of pine marten on

stumps or on forest paths

sausage-like

in
it

form, length 8-10

is

cm

with diameter about 1-1.5 cm. Usually,

spirally twisted.

Other than its dimensions, it is well distinguished by remains of small animals and birds (hair, bones, feathers), together with fruits and insects, and by the very palpable odor of musk. Sometimes, one may see the double footprints of the marten track on moist

ground or on mud. These tracks are larger and more elongate than
those of the forest polecat.

On snow,
to the
ties

the tracks of pine marten are not clearly printed due

dense fur on the food pads; in the stone marten, the callosiof the feet are always obvious in the footprints, even in winter. The most typical gait of the pine marten is a bound, leaving on
footprints.

the

snow an extended chain of paired

Sometimes, the pine

marten "trots" and

in this case, its tracks are similar to those of hares.

The pine martens rarely walks, putting one foot behind the other. The length of the bound of a marten is 65-70 cm, and on ascending a slope, not more than 45-50 cm; when ambushing its prey, up to 35-40 cm. The size of the individual prints, and the length of leaps in females, are less than in males. The animal's sex
by the relation of urine spots to the path of the tracks (during one hunting course, the marten urinates 7-8 times; Nasimovich, 1948). The marten sinks no deeper than 3-6 cm into snow. On jumpis

easily determined

ing

down from

a tree, the marten's tail does not leave a

mark

as

the sable's does (P. Yu.).

Practical Significance
In value to the fur trade, the skin of the pine marten
is
is

exceeded
durable,

only by that of sea

otter,

beaver, and

sable. Its fur

warm, and

beautiful.

Due

to its relatively small population

and the

873

584

Fig. 215. Tracks of pine marten in short (130-150 cm*) leaps; the footprints are arranged in pairs and in fours. Talitsy, Moscow district. 18 June 1951. Sketch by A.N. Formozov, about 1/5 natural size.

difficulty of capture, the pine

marten does not belong to the ob-

jects of

mass exploitation. The greatest proportion of martens is hunted in the montane forests of the northern Caucasus. Here, the most valuable and larger martens are obtained. The trade of pine marten in the USSR constitutes about 25% of the world catch of
martens (including stone marten).
the necessary prohibition of

Exhaustion of their stock as a result of overhunting has led to its hunting everywhere. This measure
to

was shown

be very effective and the prohibition was replaced

* Inconsistent with figures given in text

Sci. Ed.

874

by licenced hunting, a measure which proved

to

be the best form

of exploitation. In recent years, the stock of pine marten has not 585 been fully exploited due to insufficiently experienced hunters and

good hunting dogs. The main methods for commercial taking of marten
1)

are:

shooting with the aid of hounds; 2) driving with the aid of

hounds; 3) log deadfall traps (of the kulemok type "torlo" "snettsi", etc.*; 4) metal jaw traps. The active method of harvest with the aid
is the most effective one, but for this, good dogs are The period of harvest must be restricted to two months: December-January. The effectiveness of the various methods of

of a dog,
essential.

capture for pine marten

is

only

known

for the northwestern

is

Caucacus
a laika

(Ryabov, 1958). Tracking after the first snowfall

in

most

effective, especially in deciduous forests.

dog comes

Hunting with the aid of second place; harvest by snaring is third;

fourth

using log deadfall traps and fifth metal jaw traps. The average daily income of a hunter ranges from 8 roubles 12 kopecks to 3 roubles 18 kopecks depending upon the method of hunting.

Since the pine marten can adapt

itself to cultivated

landscapes

and lives in small forest islands of few a hundred hectares, it can be a prospective animal in the hunting trade. With shelterbelts attaining mature growth marten would be able to establish themselves widely.

With the regulation of harvest, the number of pine marten is and stable level. Between 1926-1928 (Danilov, 1963), the catch of marten constituted about 1.8% of all furs obtained in the USSR. From 1956-1959, it comprised 3.4%. The demand for skins of pine marten in the world market is not stable and is determined by styles in fur (P.Yu.).
easily maintained at a high

STONE MARTEN38
Martes (Martes) foina Erxleben,** 1777
1911 Mustela foina. Erxleben, Syst. Regni Anim.,
.

1,

p.

458.

Germany.

Special
and among

sorts of traps

Sci. Ed.

^^Also, belodushka (white-breast), " kunitsa-belodushka" (white-breasted marten)

furriers, kunitsa

Misprinted

in small lower-case type in

gorskaya (mountain marten). Russian original

Sci. Ed.

875
1192. Mustela domestica. Pinel. Actes Soc. H.N. Paris,
France.
\?>Q\.
p. 55.

1,

Mustela

foina
1,

alba.
p.

Bechstein.

Gemeinn.

Naturg.

Deutschlands,

759. Thuringia, Germany.

1869. Mustela martes var. fagorum. Fatio. Faune vertebr. Suisse,

1,

p.

318. Switzerland.
antrop.

1873. Mustela intermedia. Severtzov. Vertik. gorizont. raspredel.

Turkestansk. zhivotnykh. Izv. Obshch. lyubit.

etnogr., 8,

est.,

No.

2, p. 61.

Karagodzhur

in the

sources of the

Chu

river south* of Issyk-Kul'.

\S79. Mustela leucolachnea. Blanford. Second Yarkand Mission, Mammalia, p. 26. Yarkand, northwestern China. 1906. Mustela foina nehringi. Satunin. Izv. Kavk. muzeya,
120, tab. 2. Tbilisi.
2, p.

Transcaucasus.

1911. Martes foina bosniaca. Brass.

Aus dem Reiche der

Pelze, p.

468. Bosnia, Yugoslavia. 1914. Martes foina altaica. Satunin. Opredel. Mlekopit. Ross. Imp.

key for mammals of the Russian Empire.

estestvoisp. lyubit. prirody, 7,
1

1, p.

111. Altai.

1917. Martes rosanovi. V. et E. Martino. Zap. Krymsk. obshch.

(reprint).

Northwestern slope

of Chatyr-Dag, Crimea (Crimean preserve; V.H.).

1946. Martes foina ognevi. Laptev. Izv. Turkm.
p. 57.
fil.

AN

SSSR,

2,

Central Kopet-Dag, Bol'shoi Balkhan (type and type

It is

locality not indicated).

assumed

that the type locality is

the central Kopet-Dag; (V.H.).

586

Diagnosis
Length of
tail

with terminal hairs somewhat more than half of

body

monotone, on throat and chest a sharply outlined patch of pure white color, varies in form and size, but which usually forms two projections directed backwards to base of forelegs. Head not lighter than back. Bony tympanic bullae
length. Color

relatively short

and widely separated. Longitudinal diameter of the

is

inner half of the upper molar

half (V.H.).

only a

little

larger than the outer

The river

is

west of Issyk-Kul'

Sci. Ed.

876

586 Fig. 216. Stone marten, or white-breast, Martes (Martes) foina Erx!. Sketch by A.N. Kamarov.

Description

The stone marten

is

similar, in

its

general appearance and

It

many

individual features, to the pine marten.

has, however, on average

a somewhat longer tail, the head is more elongated, as if somewhat compressed laterally, anteriorly more pointed and, on the whole, not so rounded as that of the pine marten. Moreover, the ears of the stone marten are shorter and with more rounded tips. They are also widely separated (in the pine marten, the distance between 54-56 mm; their inner edges is about 47 mm, in the stone marten

Schmidt, 1943). In stone marten, the bare

tip

of the nose

is light,
it

usually of light flesh-color or grayish; in the pine marten,

is

dark-black or grayish-black (Schmidt, 1943). All these characteristics

make

the head

form and "face" of each marten species quite

different.

877

The

feet of the stone

marten are less furry than those of the

pine marten, and do not look as broad. Even in winter fur the heel

pads are obvious, as well as the digital. In summer fur, the legs are less fur covered, and the tail appears still longer; the difference in
ear length
in a
is more noticeable. The stone marten stands and moves manner which differs considerably from that of the pine mar-

ten

they often
is

appear to be "creeping" like the polecat, and do

not bound ("gallop") as do the pine marten and sable. This, evidently, depends also on the fact that the length of the forelegs

(height at the "withers") differs in each species

ten,
it

in the pine

mar-

about 15 cm, and in the stone marten

about

12

cm

(Schmidt, 1943).

The pelage is coarser than that of the pine marten, with elastic guard hairs and less dense underfur. The summer fur is short, sparse and coarse, and the tail is lightly furred.
The general color tone of the fur approaches that of the pine not is somewhat lighter. The underfur is also lighter color of the back dark-brown; the grayish, but whitish. The tail is
marten, but

587 is darker than that of the pine marten. In the remainder, its color
is

like that of latter.

The differences

in color

species are very sharp. In the stone marten,

and form of the throat patch of both it is always white, and

it

only in the form of a rare exception has

large and, as a rule,

light pale highlights. It

588

two projections extend backwards to the is base of the forelegs and also extends upward on the legs, sometimes halfway. Thus, the dark color of the belly juts out between the forelegs as a line into the white color of the chest and sometimes into the neck. In the pine martens, on the contrary, the white color between the forelegs juts backwards as a protrusion into the
belly color.

Numerous and various deviations from the described typical form of the patch occur, making it highly variable. Thus, the patch

may have

the shape of a bracket lacking the large anterior field, or sometimes dark spots are found in it, breaking down its form and greatly reducing its area. The patch is sometimes almost unnoticeable or absent (mainly in martens of Crete and Middle Asia). Finally, very large patches occur, of a more or less rounded form and not divided from behind. As indicated above, the form and

color of the patch are also variable in the pine marten. Variation
in this character in

both species gives, in some cases, similar forms

878

587

Fig. 217. Variations in form of throat patch of stone marten, M. (M.) foina Erxl. Upper row European marten, M. (M.)f. foina Erxl. (Middle Europe); middle row ^first and third from left Caucasian stone marten, M. (M.) / nehringi Sat. (Vladikavkaz and Talysh), second and fourth from the left Crimean stone marten, M. (M.) / rosanovi V. et E. Mart. (Crimean preserve); lower row Middle Asia stone marten, M. (M.) f. intermedia Sev. (Kopet-Dag). Drawings by N.N. Kondakov, after material of Zoological Museum,

Moscow
*Not

University (upper row, after Wehrli, 1932*, with modifications).

in Lit.

Cit. Sci. Ed.

879
of the patch. The use of this feature alone

may

lead to mistakes in

identification and to mistaken concepts of distribution, especially

of the stone marten (see below, "Geographic Distribution") and to an ungrounded confirmation concerning hybridization between the
species.

On

the whole, individual and geographic variations in color

(not considering the patch) and fur quality of stone marten are less

than in pine marten. In particular, there are not great changes in

general color tone, and no tendency towards geographic localization

is

observed.
is

There
fur.

no sexual differences
in

Young animals

their first

and character of the have autumn fur not differing

in color

from the coat of adults. The skull of the white-breast is similar to that of the pine marten and differs from it chiefly in the following features: 1. Facial portion shortened (distance from posterior edge of suborbital opening to posterior edge of canine alveolus equal to or a little more than half of distance between ends of supraorbital processes). 2. In upper profile, facial portion of skull pulled down;
therefore, convexity

formed

in

interorbital region (frontal proc-

ess
3.

feature better developed in adults and older individuals).

middle part

Nasal bones have a well-marked constriction ("isthmus") in their (a character which is noticeable mainly in younger

animals). 4. Constriction ("isthmus") on skull behind supraorbital

process more sharply defined (lines, limiting skull in this region

5. Bony tympanic bullae somewhat shorter and somewhat more widely separated, especially at their posterior part (length of bulla less, rarely equal to, distance between them at

form an angle).

their

notch absent or weakly defined.

edge of sphenopalatine carnassial tooth somethan diameter of upper molar more what larger (its length usually lying next to it). 8. General dimensions of upper molar relatively
middle
part). 6. Projection at anterior
7.

Upper

smaller and

its

inner blade considerably smaller than in pine

is

marten. In linear measurements, this blade

only a
it.

bit larger

than

the outer one, rarely equal or almost equal to

Its

longitudinal
9.

diameter

is

considerably less than diameter of entire tooth.

On
is

outer (lateral or posterolateral) surface of upper molar, there

clearly defined vertical fissure (this tooth surface
in pine marten).
is

evenly convex

Characteristics of structure of upper molar are

noteworthy for differentiation of skulls of the two marten species

(Fig. 190).

880

589

Fig. 218. Skull of stone marten,

Maries (Maries) foina

Erxl.

Sexual and age differences in the skull are the same as those
given above for the pine marten.

The number of caudal vertebrae

is

20-22 (Caucasus),

i.e.

as in

pine marten (Yurgenson, 1956), although only 17-19 are usually

recorded (Schmidt, 1943). Thorasic vertebrae are 14, lumbar,

pelvic, 3. Clavicles are smaller than in pine marten; their length

6,
is

about 10
all

mm. As

in

pine marten, they are flattened, but they are

elongated, of identical width throughout their whole length and

evenly curved (Shtreili, 1932).*

Differences in the structure of the male genital organs of both
species are quite significant.
590 larger

The penis of
about 33

the stone marten

(Shtreili,

is

its

preputial part
is

is

mm

1932).

The
its
is

form of the os penis

the same, but larger and massive in

dimensions (see below); the os penis of a young stone marten

somewhat longer than that of an old pine marten. Moreover, the generally weak sigmoid flexure of the bone is somewhat more sharply displayed. In young martens, the bone is not only shorter,
but also not thickened in
its basal part, which character is wellmarked in the adults. The diploid number of chromosomes (2N) is 38 (Vorontsov, 1958). The average dimensions of the stone marten are a little less

than the pine marten; however, this difference

is

very insignificant.

Within the species (including Central European martens), dimensions are as follows (Ognev, 1931; Kuznetsov, 1941, Schmidt,
1943; Ryabov, 1958). Body length of males 430-590, of females 380-470 mm; tail length of males 250-320, of females 230-275 mm; length of hind foot of males 85-95 mm; height of ears of males 43-47 mm; height at shoulder about 120 mm.

73.2-82.8

Condylobasal length of skull of males 71.0-86.0; of females mm; zygomatic width of males 42.2-56.4, of females,

43.6 to 50.6
29.1 to 31.0

height of skull of males 30.1-32.6, of females, (measurements of skull after data of Kuznetsov, 1941, 31 males, 29 females). Weight of males in winter is 1700-1800, in summer the aver-

mm;

mm

2000-2100 and not more than 2400 gm, that of females in winter is 1100-1300, in summer 1400-1500 gm (German martens on farm; Schmidt, 1943). For Caucasian martens, it was shown (Ryabov, 1958) that weight of males in winter was 1070-M 13251950 gm, of females 865-M 1060 gm-1306 gm (20 specimens in
age
is

*Not

in Lit.

Cit. Sci. Ed.

882
all).

These data, however, are very approximate, because they were

obtained from skinned carcasses, "in a non-fresh form or after a

in formalin." Weight of the fresh hide of a male (1) is 285 gm. Length of the os penis of young martens (15 juveniles) is 50.654.8-58.2 mm; weight is 220-320 mg. In animals two years old and more (14), length of the bone is 56.0-M 60.0-66.2 mm, weight is 380-800 mg (Caucasian martens; Ryabov, 1958). Dimensions of females are somewhat smaller than those of

long time

males. In body length, this difference

100

mm

(in the Central

is not, however, more than European martens). Dimensions are sub-

ject to negligible geographic variations.

On

the whole, variation in

meristic characters within the

in the pine

USSR,

as in color, is less than that

marten (V.H.).

Systematic Position

The stone marten

is

a well-defined species. Regardless of signifiit

cant similarity of features,

is

relatively distant

from the pine

marten. In any case, these two species are more different from

each other, than are the sable and pine marten. All "small" marten; i.e. all species of the genus Martes, except the kharza [yellowthroat marten] (M. flavigula) and il'ka [fisher] (M. pennanti), are
clearly divided into

and

all

the remainder;
if

americana), even,
It

the stone marten constitutes one and American martens (M. they are considered separate species the
two groups
i.e.

sable, pine

other.

is

characteristic that, although in the greater part of the

range of the pine marten they are encountered together (Caucasus,

central

and western Europe), hybrids between them are unknown, showing the generic relationships of both species. Scattered information about such hybridization, which appeared and continues to appear in the literature, has not received any confirmation. They are, apparently, based on specimens with throat patches, combining the color characters of one species and of the form of another. For example, the white patch in a form typical of the pine marten, or a yellow (yellowish) tone of the form which is characteristic of the stone marten. With the great variation in form of the patch in both species, and of its color in the pine marten, such combinations are encountered. In such cases, accurate identification
is

only

possible from the skull.

883

591

spring,

Only one, not fully defined, case is known, of a hybrid offwhich died immediately after its birth on a farm (Shtreili, 1932). Moreover, there is no indication about any sort of hybrid

population analogous to the kidas. Confirmation of the possibility

of appearance of individual hybrid animals (Shtreili, 1932 and

others) are without foundation.

One

of the reasons for the absence

of hybridization could be supported by essential differences in the

structure of the
in

male sex organs (see above

is

in

corresponding places
to the

"Description" sections of both species).

The stone marten

pine marten.
It is,

more ancient species compared

their habits

apparently, a Pliocene form. Both species are

strongly differentiated from each other by ogy (Schmidt, 1943) (V.H.).

and ethol-

Geographic Distribution

Found

in

montane regions of the Near and Middle East and

Central Asia, and montane and in part plains, regions of Europe.

Geographic Range

in the Soviet

Union

Not extensive, constituting less than half the range of the species, and connected with the extreme west of the country and with its southern mountainous borders. Within the USSR, it is divided into several parts, isolated from each other, but united beyond our
borders.

The

largest

western

part of the range,

which

is

connected

with the western European part of the species range, occupies the
Baltic region and includes

Sarema [Saaremaa] Island (EzeF [Oesel]) (Ognev, 1931; Kalninysh, 1950), Lithuania, parts of Byelorussia and Ukraine and several middle Russian districts. The boundaries
In the north, the

of this part of the range are not well-understood and reliable data

on them are scarce.

boundary

starts at the

Gulf of

Finland west of Leningrad, apparently somewhere on the meridian

of Chudsk Lake (the marten
Lithuania.
is known from Rakvera, the former Vezenberg), and extend southward from western Estonia, and

It bypasses Leningrad and Pskovsk districts on the west and passes somewhere along the middle part of Byelorussia. For this part of the USSR, its range has been shown to include Grodnensk, Minsk, Brest and Gomel'sk districts (Serzhanin, 1955,

1961).

884
Farther, the

boundary turns again,

to the east in a not yet

accurately established line including southern Byelorussia and

Poles'ia (Gomel'sk district, Mozyr', Fedyushin, 1928; Serzhanin,
1956), passes eastward, occupying the southwestern part of Bryansk
district

data of A.V. Fedesov

this species
It

Surazha, Pochepa, Trubchevska, and Pogara regions (new but according Fedosov and 1951,
to

Nikitin,

is

not recorded in Bryansk district; Melander, 1938).

is

not excluded that this marten lives in the southern parts of

district.

Mogilevsk

From

the indicated places in Bryansk district, the range bound-

ary goes on to NovosiF (east of Orel, former Tula governance; Ognev, 1931). In the lower Svapa, the boundary lies to the south

of this line, south of Dmitriev city (Ptushenko, 1937).

From

Novosil,

boundary passes to Bobrov (Sadovoe; Ognev and Vorob'ev, stone marten was recorded in Chernaya Kalitva (Budenovsk and
the

region southeast of Voronezh

1924). South of Voronezh, the
the

upper Tikhaya-Sosna and Ladomirovsk regions; BarabashNikiforov, 1957). From Bobrov, the boundary is directed to the south and then to the west in a not yet exactly established line,
forming the southern border of this part of the range. It proceeds north of Lugansk, including the region to the northwest, then descends to Donetsk, and proceeds farther, somewhere in the
Melitopol region or somewhat to the north, extending to Perekop
or to Sivash (after Korneev, 1952). Farther to the west, the Black

Sea coast forms the border.

Note. The outline of the northern and eastern boundaries of the

western part of the range differs greatly from those usually presented in reports. Thus,

Ognev (1931, 1947) records

the existence

of the stone marten in "former Pskov, Vitebsk and Smolensk

governances", Yaroslavl' and Kaluga districts and believes in

"established" occurrence in

its

and "most probably" in the former Pereslavsk co[unty] of Vladimir governance and so on. Yurgenson (1932, 1933) describes the border of the range from
district

Moscow

Chudsk Lake
and nearly
the
593

to the

upper Volga, and along

it

[river] to Zhigulei

to Saratov (map),

and then to Dnepropetrovsk and along

are
a

Dnepr
In

to the

Black Sea.
there
series

the

literature

of precisely

known
This

occurrences of stone marten outside the above-outlined

line

line.

was

definitely or indefinitely noted for former Yaroslavsk gov-

ernance and Yaroslavl,

Moscow

district,

Ivanovsk

district,

Zhigulei

885

and even for the north and middle Urals, Kaslinsk Urals and Pechora-Ilych Territory (Sabaneev, 1868, 1872, 1874; Bogdanov, 1873; Varentsov, 1919, Shillinger, 1929; Ognev, 1931, 1947;
Yurgenson, 1932, 1933).

Some

of this information

is

simply fantastic. Such are data

given by Sabaneev,

known

for his errors, for the Urals or about

Pechora-Ilych

Territory;

information

about

Yaroslavl

and

Yaroslavsk oblast, also given by Sabaneev, did not receive any confirmation. These martens were not even observed in the Valdai
Hills (Stroganov, 1934, 1936). There are

no

facts suggesting their

existence in the former Pereslavsk county of Vladimir governance

and

in

the Ivanovsk district and especially, in Zhigulya^^ and

farther

down along

the

Volga almost

to Saratov. All of this infor-

mation should be rejected, although on some of them, the most dubious (Urals), historical-geographical hypotheses were based.

Assumptions concerning the former Pskovsk and Vitebsk

if these areas are considered as corresponding to the contemporary districts, are interpreted as follows. The most westerly part of former Vitebsk governance is included in Lithuania, and the stone marten is recorded in this republic. Concerning former Pskovsk governance, in the extreme western part, which coincides with the west of present Pskovsk district, transgressions or

governances,

sporadic inhabitation are possible.

the range actually passes along the border of this district.

existence of the
established.

The above described border of The described species in Smolensk district still is not
be given
dis-

From
trict,

all

that has preceded, particular attention should

to information

on occurrence of the stone marten

southern outskirts and
at its border.

in

Moscow

along

its

Old and poorly

at that

defined records (Sabaneev, 1872) for the district

period

were disproved (Satunin, 1892). At the beginning and middle of the 20th century, however, newer data appeared at the Taratin quarries, near Maloyaroslavets (former Kaluga governance hence the reference to this governance), the lake between Lukhovitsi and Stupoe, and Shchurov (Shchurov quarries) near Golutvin (Ognev, 1931, 1947; Troshin, 1961). Not all of these records were

''

Bogdanov (1873) wrote: "Until now

It

have not succeeded

in learning

anything
area;

positive about the existence of the marten, M. foina Briss. with us (middle

Volga

V.H.).

is

highly probable that

it

is

present in "Zhigulya". Later, in literature, this

careful assumption

became

the precedence for serious confirmation.

887

confirmed by documented scientific material, and

to consider

it is
it

not possible

them confirmed. However, neither can

be fully ex-

and sporadically be found had not been recorded there Novosil' in the former Tula governance now lies in the present Orlov district) and, from here, individual animals may penetrate northwards to Oka and beyond. It is possible, that they dwell permanently somewhere here. Captures of stone marten in Ozeri is
rarely

cluded that the stone marten

may

somewhere

in the

Tula

district

(it

connected with

its

penetration thence from

Ryazan

district (Troshin,

1961), where marten was introduced for acclimatization (see

below). There are no objections to this assumption.

The

prior erroneous information

was based

either on simple

mistakes (capture of pine martens with pure white patch) or on

information given by poorly informed people led astray by that
character. Identification of a specimen obtained
eral part of the

from the periphto

range absolutely should also be based on skull.

Insufficiently critical consideration of reports

on the genus led

misrepresentations in revisions (Ognev, 1931; Novikov, 1956 and

particularly Yurgenson, 1933).

Some

corrections to the boundary of the range of marten are

is reliable.

possible, but at the present time, only that outlined above

In the Crimea, the marten
is

widely distributed
is

in the

moun-

tainous part of the peninsula, and

plains at a distance

apparently not found in the

from the mountains. Thus, the Crimean section

of the range

is,

apparently, separated from the northern part.

The Caucasian section of the range is isolated both from the Crimean and the western parts. Its northern border passes from the lower Kuban, at first along the northern foothills of the Main Range,
then proceeding as a large projection northward, including
all

the

Stavropol uplands, and along the foothills again, extends to

Makhachkala;
pian coast,
it

i.e.

the shore of the Caspian Sea.

Along the valley

of the Terek, the marten extends east of Grozny. Along the Cas-

reaches or almost reaches the sea through the gallery

Samur [river] as far as its mouth; it is encountered on the Apsheron peninsula (Heptner and Formozov, 1941;
forests of the

Vereshchagin, 1947, 1959). In the south, the range extends to the

state

boundary and beyond

it.

The marten

is

absent only in the

desert plains of the eastern Transcaucasus (Vereshchagin, 1947,

1959), although, according to older data,

(Satunin, 1915).

it

was met with there

888
594

The stone marten

within the
sian range.

is

present in Talysh. This area of occurrence

USSR

is,

possibly, isolated

from the

rest of the

Cauca-

The Middle Asian and southern Siberian parts of the range in our country are divided into a number of separate parts. In the west, the marten exists throughout the Kopet-Dag [range] down to
its foot,

along the Atrek

[river], at least its

upper course, and in the

(right

Bol'shoi Balkhan [range] (Laptev, 1934; V.G. Heptner). In southern Turkmenia,

it is

encountered

in

Gyaz'-Gyadyk

bank of

upper Tedzhen, V.G. Heptner) and in the mountainous region between Kushka and the upper Murgab (Flerov, 1932; V.G.
Heptner). These are extensions of the range coming from Iran and

Afghanistan.
large and includes
to

beyond the Amu-Dar'ya is quite from the Pamiro-Alaisk the Tien Shan systems. Westward the range extends apparently

The

section of the range lying

all

the mountain ranges

along the ranges to the western extremities of the Gissar (V.G.

Heptner), Zeravashan, Turkestan ranges and even to the Nuratau

(Meklenburtsev, 1937).

To

the north,

it

occupies the Karatau, Talas

and Kirgiz ranges and the Zailiisk Alatau, including its western spurs. Within the range are included also the northern spurs of this range, the Dzhungarsk Alatau and the small Kzyl-Togai mountains in Alakul' depression (Sludskii, 1953). The stone marten is only
absent in the Eastern Pamir, Farther to the east, a section of the
595

range occupies the Tarbagatai and Saur [mountains].

its

It

also inhab-

the Kalbinsk Altai (Kuznetsov, 1948). All of these indicated

places are connected with

Afghan and Central Asian regions

in-

habited by the stone marten.

A
Ul'ba

separate part of the range occupies southern Altai.

is

ten here

distributed southward to the

[rivers] (Sludskii,

The marKurchum, Bukhtarma and 1953), and eastward to the frontier. The

northern distributional limits in the Altai are not clear as to the

accurate limits of distribution to the east. Apparently, the marten
is

absent

at Telets

Lake and along

is

the

Chulyshman (Yurgenson,

1938; V.G. Heptner). In the Sayan and in Tuva, particularly in

Tanna-Ola, the marten

absent (Yanushevich, 1952).

The stone marten was introduced (59 animals in 1936) on the right bank of the Oka above the mouth of the Moksha (Ryazan
district;

Lavrov, 1946). This measure, naturally, failed to succeed;

the marten, for 25 years, did not

become

commercial species;

889

594

Fig. 220. Species range of the stone marten,

Martes (Martes) foina Erxl. (V.G. Heptner)

however, a small number of animals were, apparently, preserved

in

Ryazan

They spread through the northern forest past of the district, from where they had been introduced for a distance of about 100 km. From 1936 to 1952, individual cases of marten capture were known in the Kasimov region (near the place of inoblast.

troduction), in Gusya-Zheleznyi and

Tuma regions, and

near Spassk

(Nazarov, 1957).

890

Geographic Range Outside the Soviet Union

This occupies southern and central Europe from the Mediterranean Sea westward to the Atlantic Ocean (absent in England and Ireland) and northwards to the Baltic coast, including

Denmark and

some

islands in the Mediterranean Sea (Rhodes, Crete [Corfu];

absent on the Balearics, Sardinia, Corsica and Sicily. In Asia, the range includes Asia Minor, Syria, Palestine, montane parts of Iraq,

northern and western parts of Iran, Afghanistan, Baluchistan, Chitral, Kashmir (northern), northern (montane) Punjab, Nepal,
Sikkim); in China
the region
Tibet,'^

probably Shansi and Chihli (Hebei),

i.e.

between the Huang He, the ocean and the Great Wall.
it

This does not exclude the possibility that

may

penetrate to the

extreme southern part of northeastern China (see below). In the north, the range is a narrow extension isolated from the previously-described Central Asiatic region, stretching eastward

and occupying the montane parts of eastern Kashgariya (Tien Shan system), the western montane part of Dzhungariya (rising up to connect with the Dzhungarsk Alatau and Tarbagatai) and the northwestern and northern parts of the Mongolian Republic. Here, the range occupies, it seems, the Mongolian Altai southeastward,

approximately to 100 (Ikhe-Bogdo), the Kobdo region, PriKosogoF

mountains, and perhaps the Khangai.

one sometimes encounters information on the occurrence of the stone marten in northeastern China (former Manchuria; Bobrinskii, 1944 and others). This information is evidently wrong. The fact is that in the USSR, the stone marten
In the literature

was never recorded anywhere east of the southern Altai, or in the Mongolian Republic except where mentioned above, where it is very rare; it is not recorded in Kentei. Old information about the occurrence of the stone marten in the Mongolian Republic in the Great Khingan [mountains] (Borodovskii, 1894) and considered
correct,
is false (Bannikov, 1954). Concerning northeastern China, there are no accurate and reliable data about occurrence here of any species of the genus Martes, except sable (series of papers, particularly in 1934 by Lukashkin;

""No details on distribution

1931).

in

Tibet are known.

is

It

is

very probable that

not occupy the whole country. There

information for

Kam

it does and Nan-Shan (Ognev,

The

basis for confirmation of

its

occurrence in the Nan-Shan

is

not clear.

Reported for Lhasa.

891

also in the Great

Lukashkin and Zhernkov, 1934). The reports of Sowerby (1923), whose mistakes are

Khingan according
reliable'*'

to

recognized, are not

(see also range description of pine

marten, page 846), or

may be based on
is

individual variants of

Man-

churian sable, the quality of which

significantly poorer than that

of Siberian.
Nevertheless, the possibility that marten

may appear

in

the

extreme south of northeastern China in regions adjoining northern

China

is

not excluded, although very doubtful. However, their very

In

presence here (Shansi and Chihli-Hebei) calls for skepticism*.

any case,

it

is

now

generally believed that in northeastern China,

northern (Great Khingan, Il'khuri-Alin') and eastern Manchurian mountains) martens are absent. The whole question of the existence of stone marten in China demands speat least in its

parts (eastern

cial analysis.

596

Information distributed in our literature about

its

occurrence in

southern Finland (Ognev, 1931; Yurgenson, 1932; Novikov, 1956)

is

mistaken. The stone marten was nowhere in this country

(Siivonen, 1956;

Van den

Brink, 1958).
is

In the zoogeographical literature, the stone marten

usually
the

evaluated as "European", or "western" form.

outlined range,
part a
it is

As seen from

typically and Near- and Central Asiatic, and in

"Western Mediterranean" species closely associated with

it

mountains. Only, secondarily, apparently, did

Its

settle in

Europe.

occurrence in the European plain far from the mountains, but

is

not in other places,

apparently connected to a significant extent

it

with humans and their constructions. In Central Europe,

is

well-marked synanthropic species (V.H.).

Geographic Variation
Geographic variation of the stone marten within the USSR is insignificant. This is related to the amplitude of racial differences, as
well as the

number of forms.

It is

highly probable, that their

number

the country (former Manchuria; V.H.),

abundant in the western portion of range spreads westwards into the mountains of eastern Mongolia and on through northern Chili, Shansi and into West China. I have seen large consignments of skins, from Mukden, as well as from North Shansi, and can detect no difference between the specimens from the two regions" (p.

"This author, for example,

states that "It is

whence

its

68)**.

*Presence confirmed by Zhang

**In the copy
I

et al.,
this

1997

is

Sci. Ed.

have consulted,

quote

on

p.

66, paragraph 3

Sci. Ed.

892

what is usually considered. The fur industry differentiates only two groups of stone marten: the Middle Asian, to which the martens of Middle Asia belong; and the Caucasian to which the martens of the European part of the Union and Caucasus beis

less than

long. Evidently, geographic variation in that part of the range

lies

which

outside our borders

is

also overestimated.

The

entire question

needs to be reexamined with solid material.

For the

territory of our country, four

forms are usually consid-

ered (chiefly based on data of Kuznetsov, 1941, and also

Ognev

1931; Shtreili, 1932*; Schmidt, 1943; Ryabov, 1958 and others

with modifications).
1.

European stone marten M. {M.)f.foina Erxleben, 1777

size.

(syn.

domestica, alba, fagorum).

General dimensions quite small, skull of average

In winter fur, color of

tawny

to

back varies from quite light grayish completely dark brown. Guard hairs are tawny or chestis

nut-brown, underfur
lighter than

very

light,

pale-gray or whitish, sides a bit

back and belly darker. Withers slightly lighter than back. Tail dark, brown or blackish-brown in color, darker at end.

Legs dark brown. Throat patch pure white, variable

form.

in size

and

Body

length of adult males usually

Tail

400-440 mm. 250 mm.

84.6

450-500 mm; of females, length of males 250-270 mm, of females 230-

Condylobasal length of male skull (9) 71.0-M 82.00 0.65mm; of females (5) 77.6-M 78.8-80.0 mm; zygomatic width of males 47.9-M 51.7 0.75-52.8 mm, of females 46.2-M 47.850.0 mm; height of male skull 31.0-M 31.9 0.49-32.6 mm.
Present in European part of the

Outside the

USSR in

USSR, except Crimea.

western Europe, except the Balkan

Peninsula and Pyrenees [Iberian] Peninsula.

the nominal form, so far as

mentioned belongs to known. Material of this form from our country, and from actual Central Europe, is very poor, and a special comparison of the East and Central European martens at a modern level has not been done. Nevertheless, the identity of these
the part of our country
is

The marten of

populations

is

sufficiently obvious.

Descriptions of fur and dimensions are based on martens of

Central Europe.

Not

in Lit.

Cit. Sci. Ed.

893

2.

Crimean stone marten, M. (M.) f. rosanovi V.

in

et E.

Martino,

1917.

Dimensions somewhat smaller than

small, not massive.
597

preceding form. Skull

Color of winter fur near or identical with the color of previous

form. Back quite dark, smoky-tawny in color, with dark-tawny

guard hairs and whitish underfur. Throat patch variable in form

and
tail

size.

(1) 470 mm; mm, of females 275 mm, length of hind foot of males 94-95 mm; height of ear of males 43-47 mm, 41 mm. of females Condylobasal length of male skull (4) M 78 mm, of females (5) 73.2-M 75.0-76.9 mm; zygomatic width of males 42.8-M 43.848.1 mm, of females 42.6-M 45.7-47.0 mm.

Body

length of males

465-510 mm, of females

length of males 300-325

In

montane Crimea.

Outside the

USSR,

absent.
is

The Crimean form

very near to that described above, and

it

is,

apparently, identical with

(Ognev, 1931, equated the southern

Ukraine marten with the Crimean form). The only difference lies in the somewhat smaller average dimensions of the Crimean
martens. Beside, this was established on insufficient material. Other
features given earlier

(some

particularities in tooth structure;

Martino, 1917; Ognev, 1931) are not well-founded. The question

requires re-examination.
3. Caucasian stone marten, M. (M.) f. nehringi Satunin, 1905. Dimensions large this form is, apparently, the largest in the species within the USSR. Skull large and massive. Winter coat quite dark, brownish-tawny or dark tawny with grayish tint, depending on effect of light-grayish or whitish underfur. Sides lighter than back, because here guard hairs are fewer and light underfur shines through more strongly. Tail and feet dark brown. Throat patch very variable in form and size, shows a tendency towards reduction. Body length about 540 mm, tail length 250-260 mm, length of hind feet 90-100 mm. Condylobasal length of male skulH^ (13) 81.0-M 83.9 0.29

(84.3)-86.0

mm,

of females (15)

76.1-M 79.7 0.69 (79.2)-82.3

parentheses after

^Measurements of skull, as in other cases, given after Kuznetsov (1941), figures in average of 20 specimens from Caucasian preserve (Ryabov, 1958).

894

mm; zygomatic width of the male skull 50.3-M 52.9 0.47 (51.0)mm, of females 46.2, M 48.6 0.40 (44.9)-50.6 mm; height of male skull 30.1-M 32.1 0.26 mm (31.7 mm)-33.1 mm, of
56.4 30.4 mm). (28.9 females 29.1-M 29.7 0.14 For dimensions of os penis, see above, "Description" section.
In Caucasus.

mm

mm

Outside the
Iran and Turkey.

USSR, probably

present in contiguous parts of

Differences between this form and the nominal are not sharp.

Moreover,

it is

not clear whether the marten of the

Main range and

those of the southern xeric part of Trans-Caucasus are identical.

There are signs of similarity between Caucasian martens and the Balkan form bosniaca (Kuznetsov, 1941), and of the relationship of martens of the Main range to the nominal form, and only of Trans-Caucasian, to the described form (Novikov, 1956).
4. Middle Asian stone marten, M. (M.)f. intermedia Severtzov, 1873 (syn. leucolachnea, altaica, ognevi). General dimensions, apparently, somewhat smaller than in

Caucasian form.

Color of winter fur, on average, lighter than in Caucasian marten. General tone of back grayish-tawny of moderate darkness. Sides lighter, but of same tone as back. Guard hairs dark-tawny,
598

underfur almost white,

tail

dark brown; throat patch very variable,

sometimes completely undefined. Condylobasal length of male skull (5) 81.2-M 83.2-84.6 mm, of females (4) 77.7-M77.9-78.0 mm; zygomatic width of males 49.951.4-52.0 mm, of females 46.5-M 47.0-47.4 mm; height of male

skull

30.5-M 31.6-32.5 mm, of females 29.8-M 30.3-31.0 mm. In montane Middle Asia, from Kopet-Dag and Bol'shoi Balkhan
and
Altai.
in northern Iran (?),

to Tarbagatai

Outside the

USSR,

Afghanistan, western
(?),

Pakistan, western Himalayas, Tien Shan in China, Tibet

and

northern part of Mongolian Republic.

It is

not excluded that the marten of the extreme west (Kopet-

Dag)

from the eastern (Tien Shan); however, the distinction of the form ognevi was presented without adequate argument. Identity of Himalayan and Tien Shan martens is estabdiffer also

lished (Pocock, 1941).

895
In the parts of the range lying outside the limits of the

USSR,

the following forms are usually recognized: 1) M. (M.) /. mediterranea Barrett-Hamilton, 1898 Pyrenees [Iberian] PeninYugoslavia; 3) M. (M.) sula; 2) M. (M.)/ bosniaca Brass, 1911

milleri Festa,

1914

Island

of Rhodes; 4) M. (M.)

f.

bunites

1906 Crete; 5) M. (M.)f. syriaca Nehring, 1902 Syria; 6) M. (M.) / toufoeus Hodgson, 1842 Tibet, Lhasa; 7) M. (M.) /
Bate,

kozlovi Ognev, 1931

parently are

Kam, eastern Tibet. The last two forms apsynonyms and possibly belong to intermedia (V.H.)Biology

Population. Information

abundance of stone marten is in the Caucasus and Trans-Caucasus. In Middle Asia, they are encountered more rarely, because habitats typical for them
is

scanty.

The

greatest

are not found everywhere and are unevenly distributed.

rare in the

It is

not

Crimean Peninsula (V.G. Heptner), where

it is

met with

only in the mountains. Within Ukraine, the stone marten is mainly characteristic for the territories to the west of the Dnepr, where it
primarily found in populated areas, and more rarely in forests and forest shelter-belts, and not everywhere, but sporadically. It is quite common in the forests of Belgorod district (central forestis

steppe, Novikov, 1962). In the middle [forest] zone,

rarity.

it

is

a great

Habitat.

The

habitat of the stone marten lend themselves to

division into primary and secondary.

The

latter

appear

in the plains

regions of the range where the stone marten transgresses to a significant extent into populated areas, in the role of a synanthropic

species. Here,
lars,

it

prefers to live in stone structures

in attics, cel-

sheds, stables, schools, storehouses, railway stations, in stone

walls, mill weirs,

and even
is

in old stork nests, etc.

In the central forest-steppe (Belgorod district), the stone

marten (Novikov, 1962)

a characteristic inhabitant of forests.

It

prefers old tall-trunked oak-groves on hills, especially those with

broken

relief. It

more

ests or oak-groves,

rarely settles in the middle-aged mixed forand avoid the sections with low relief. Here, it

also settles in buildings.

Primary habitats are characteristic for those parts of the range

which exhibit

more or
by
less

less distinctly

are distinguished

montane character. They harsh climate; snow cover here is not

896

permanent and
599 cally,

is

uneven. These are unforested or almost unforested

mountains, ravines, rock slides and breccias with, characteristi-

shrubby vegetation of a Mediterranean type, "maquis" or

It

sheblyak*', forests and rocky and stony

orchards, vineyards, and nut groves.

forests, but in the

mountain slopes, deserted does not avoid montane

Caucasus,

it

clearly prefers broadleaf forests in

foothills. It avoids the

dark coniferous forest zone, mainly because

of their

snowy

slides at the

winters. It is also encountered in rock outcrops and upper forest limits and even above, up to 2600-2700

above sea level. In northwestern Caucasus (Ryabov, 1959), the upper limit is only 1000-1200 m. The stone marten is not adapted living in places with continuous deep snow cover. It captures food almost exclusively on the ground, hiding in the mountains, among rocks and boulders. In heavy-snow regions, marten are deprived
of shelter when these regions are flooded in spring by the thaw. Moreover, they do not possess footpads densely covered with fur characteristic of sable and pine marten, and therefore are more
sensitive to cold.

The weight load per

marten foot
is

cm^ of

the supporting surface of the stone

double that of the pine marten (Ryabov, 1959): 30.9

this is

gm

and 15.2 gm; (Ryabov,

why

it

obliged to avoid

snowy

regions).

from abundance of plant food in its diet. In 240 feces, collected chiefly in populated places at the Pravoberezbnaya [right bank of the Dnepr] Ukraine (Pidoplichko, 1929) cherry fruits were encountered in 50%, apple and pear fruits in 25%, plums in 5%, black nightshade in 8%, tomatoes in 2.0%, mouse-like rodents only in 7%, and birds in 3%. Of 20 samples, only fruits were found in 10, and small rodents in one case. One marten ate during a winter more than 2 sacks of dry fruits in an attic. It also catches rats and mice, and rarely attacks hens. In the forests of Nikolaevsk and Dnepropetrovsk districts (296 excrements and food items), the food composition of the stone marten was as follows (Abelentsev, 1958, % of occurrence):
Food.
typical feature, distinguishing the stone marten
is

pine marten and sable,

the

Vertebrates

76.0
59.0 55.6

Plant foods

62.3

Mammals
Rodents
Birds
Russian word

Olive

fruit

21.6
fruit fruit

Blackthorn

19.9
8.5

44.6

Wild pear

for shrubby deciduous successional vegetation

Sci. Ed.

897

Sparrows
Lizards
Insects

34.1

Sunflower seed

11.8
3.5

7.0

Common

cherry fruit
fruit

50.0
50.0
4.4

Beetles
Insect larvae

Malagensk cherry Mulberry fruit Rose fruit

4.5
4.1

7.4

Among

vertebrate animals, the following were found:

Common wood mouse

Gray hamster

16.2 14.2

Hares (young)

13.8

Mole

rat

6.0
5.4 2.2

Common
In

vole

7.0
1.4

Spotted ground squirrel

Kurgan [barrow] mouse

Common
in the

hamster

summer, rodent occurrence

insect occurrences

food of martens reached

75-88%, and

75-81%.

Food composition
in

changed significantly annually, depending upon fluctuations natural abundance of various foods.

In the "Vorskl Forest" (463 specimens; majority pertain to

600

autumn and winter) (Novikov, 1962) rodent occurrence in various to 75%; in summer from 20 to years and seasons varied from to 57.9%. The 75%, in autumn from 1 1 to 33%, in winter from whole period, 7.3%, encountered: for the often vole was most red and in summer vole 0.9%, in summer, 16.1%; the subterranean 12.8%. Also found were water vole, the yellow-necked field mouse and house mouse, and field hare (in winter up to 1.5%). The occurrence of birds in the marten diet varied in summer from from to 16.7%, in winter from to 33%, in autumn 16.6% to 88.9%. These were mainly sparrow-like birds (no fewer than 17-18 species). One litter of martens was reared on the basis of leavings from a rook's nest. Marten also eat carrion. This could

partially explain the presence of poultry in

food remains (Novikov,

at-

1962). However, individual animals sometimes systematically

tack domestic fowl (Ryabov, 1959). Attacks on chicken coops by

marten

in the

Crimean preserve took place even

in the

presence of

an excess of forest mice (V.G. Heptner). For the stone marten,

birds are an attractive but difficult prey.
In summer nutrition in "Vorskl Forest", insects were usually met with in 30-66.7%. However, in the summer of 1960, they were not found at all. In autumn, their occurrence varied from 25 from to 33.3%. Among insects, beetles to 100%, in winter predominated, chiefly dung beetles captured along forest roads and paths. Overall occurrence of beetles was 22.0%.


898
Plant food (fruits) plays here an important role in autumn and
to 70%, summer, the fruits and berries occurred from from 9.17 to 100%, and in winter from to 100% of in autumn 42.7%, and apple the cases. Occurrence of sloe was 36%, pear 17.0%. No less than 27 species of plant food were revealed. The

winter. In

significance of these foods in different years fluctuates greatly.

Thus,

in the

winter of 1958, they constituted almost the only food

of the stone marten. In years poor in plant yields, plant foods were

completely absent

in the

marten's food (1947, 1962). The stone

fruits

marten feeds both on carrion, as well as

branches of trees and bushes.

hanging from the

This clearly revealed polyphagia of the stone marten usually

ensures their nutritional needs even in case of sharp fluctuations in
the yield of individual foods.

However,

in the winter

of 1947, after

a severe

summer

drought, martens were obviously hungry, feeding

on garbage, feces, etc. (Novikov, 1962). In the northwestern Caucasus, 262 yew

fruits (154 g) were one stomach, and in the rectum 43 more. In 43 excrements collected from rock slides of the Caucasian preserve, mammals were found in 85%, birds 16.3%, of which small birds were 11.6%, and invertebrates 20.2% (Donaurov, Teplov, and

found

in

Shikina, 1936).

Zakatalo-Nukhinsk valley (Azerbaidzhan), the food of the stone marten was composed of the following elements
In the

(Rukovskii, 1957;

of occurrence). 65.5
5.1

Mammals
Birds
Reptiles
Insects

Grapes

10.8
16.1

Dogwood
Myrobalan plum
Blackthorn

3.5

3.5

84.0
2.4

4.3
3.5 1.0
is

Acorns
Blackberry

Pear

23.3

Nuts

The proportion by weight of

also quite significant.
In

the insects and plant food here

montane Crimea, the stone marten eats grapes and grape According to analysis of 373 specimens in the Crimean preserve in 1936-1938 (M.I. Savvina) the food composition of the stone marten was as follows:
snails (Flervov, 1929).

Mouse-like rodents Hares

41.0
1.3

Molluscs
Pears

0.9

19.0


899

Roe deer
Birds
Lizards
Fish
Insects

(carrion?)

1.0

Dogwood
Rose Mountain ash
Juniper

7.1

16.0
0.3
0.5

9.3

0.3
0.5
1.8

50.4
9.1

Mushrooms
Conifer needles

Insect larvae

2.0

In the mountains of Kirghizia (Kuznetsov,

1948), the stone

marten eats snowcocks, partridges, raspberry and mountain ash; the latter were also observed in the mountains of Semirech'e
601

(Shnitnikov, 1936).

The old

naturalists of western
rats,

Europe (Brehm,

1866; Chudi, 1873) refer to

mice, domestic fowl, rabbits,

small birds, bird eggs, lizards, frogs, bee's honey, grapes, the fruits
of cherry, plum and pear, mountain ash and gooseberries,

hemp

seed and garden vegetables, as food items for the marten. The

phytophagous nature of

this

animal

is

here confirmed.

one day one ground squirrel weighing about 176 gm or 2 magpies. In two days, one marten ate a mole rat weighing 358 gm; when feeding on small animals about 5-8 gray hamsters (weighing 35 gm each) or 9-10 yellownecked mice, weighing 20 gm each. Thus, the daily ration was
in captivity ate in

Young martens

about 170-250
a

gm of animal food (Abelentsev, 1958). In summer, young marten ate in one day about 15 apricots whose flesh weighed about 260 gm. Upon receiving animal food, stone martens
foods were often preferred; and 250 g of apricot flesh for when offered each day 10 mice several successive days, they ate the fruits first and left the mice.

did not refuse plant food. In hot weather, they ate only mulberry
fruits, cherries, etc. In captivity, plant

The

ration for martens in the nursery [animal facility] of the

Zoological Institute of the

consisted of 150-180

Academy

of Sciences of the

gm

of rabbits, 25-30

gm

of milk, 20-30

USSR gm

cornelian cherry or briar fruits or 30

gm

boiled carrots per head.

On

At the "Vorskl Forest" a stone marten ate in an open-air cage, in one day, 2-8 small birds, and moreover, willingly ate cherries and apples. Home range. There is very little information. In one case, in the northwestern Caucasus, the area of the daily range was determined to be 400 ha and the length of the daily track was 8 km. The range was poor in food (Ryabov, 1959). In the southern Ukraine
such a ration, the martens
felt

quite satisfied.

(Abelentsev, 1958), the

maximum

length of the marten's daily track

far

was

km, but sometimes a nursing female did not go

from her

900
den. Tracing of 16 daily tracks in "Vorskl Forest" in Belgorodsk
district

showed

that the average length

was equal

to

about

km,

rarely reaching 2.5-3.0 km.

Length of the daily track depends on the abundance of food

and conditions of its availability; it is inversely proportional to the abundance and availability of food. In the starvation winter of 1946/47, martens went, in one day, over a much longer route than they did in the winter of 1957/58, when there were many small rodents and fruits (Novikov, 1962). In the first case, the average
length of the daily track was 985 m, and up to 1700-2400 m, and
in the

second case

540 m and not more than

in the

1100 m. The daily

but,

track
often,

may sometimes be
it

form of a closed loop,

more

extends in any direction. In one case, the daily range of

had an area of about 2.5 km^. Burrows and shelters. The stone marten does not dig a burrow and does not occupy foreign ones. It prefers fissures and clefts in rocks, spaces between stones in rock slides, inhabited and uninhabited stone structures, etc. It sometimes lives in tree holes at a height up to 9 m. In the forested part of the Crimean preserve, it gives birth chiefly in tree holes (V.G. Heptner). However, in the forests of the western Caucasus, of five dens, four were in rocks
activity

(Ryabov, 1959).
Daily activity and behavior. The daily activity rhythm in the marten is not exact. The marten is active mainly during twilight and at night, but cases of daytime activity are observed. It is a
crepuscular and nocturnal animal, but to a lesser extent than the

European polecat, which the marten frequently meets in one or another locality in the lowland part of its range (Shyutse, 1936)*. The marten is very active on bright moonlit nights. Being mainly
a terrestrial predator, the stone marten ten in its ability to
is

inferior to the pine mar-

However, it climbs well and in V.G. Heptner), it does so frepreserve; dense forests (Crimean quently. It is a capable swimmer. It is active by day most often in
climb
trees.

602

(Dinnik, 1914), when the nights are short. In the central forest-steppe (Belgorodsk district), the stone marten leads a

summer

crepuscular-nocturnal

way

of

life,

but in

summer

it is

observed by
in the early

day, even in midday hours. Martens living in open-air enclosures

1700-2000 hours but hid in the nest 0600-0700 hours (Novikov, 1962). morning by
were outside
at

*Not

in Lit.

Cit. Sci. Ed.

901

602

Fig. 221. Stone marten,

Martes {Maries) foina Erxl, Caucasian preserve. Photograph by L.S. Ryabov and Yu.S. Danilchenko.

The Stone marten captures its food mainly on the ground surfrom following daily tracks (Novikov, 1962) showed that along 17 km, they moved only 129 m through the tree canopy. There were 105 climbs into trees. Along the length of this route, martens examined 107 different terrestrial covers logs, brushwood
face. Result

heaps, overhanging turfs, holes, root-hollows, etc. Martens also

dug For

in the

snow 34 times

in

a considerable part of

way

open places, and near trees 23 times. stone marten went on foot, not

lingering,

their tracks are

and making only small turns. Only in thickets of shrubs, found to meander, a characteristic for the period of

fattening.

In soft snow, martens prefer to use for their

movements paths
tracks.

of hares and ski-tracks.

Martens usually move

in large

They rarely run along their own jumps measuring 30-40 cm in

or, the gait

length.

On

firm or crusted

snow

changes into small steps, and

in the track, the five foot

pads are well-defined. In summer, stone

902
martens

move

at a

walk along forest paths and roads, and

is

in the

grass, they

bound (Novikov, 1962). Seasonal migrations and transgressions. Information

absent.

Reproduction. Estrus and copulation, according to the observa-

603

same time as in pine marten. In nature, individual cases of mating were observed on 18 June and 16 July. Rut in Belgorodsk district was said to take place in June ("Vorskla Forest", Novikov, 1962). Mating occurs on the ground or on the roofs of houses. It was observed early in the morning and on moonlit nights. The period of pregnancy is also the same as in pine marten, i.e. 236-237 days (ManteifeF, 1947). Some fur breeders report 254-265 or 258-275 (in sable, average duration is 272 days). The average number of young in a litter is 3-7. In individual cases 2 young were found. Parturition takes place from the end of
tions of fur breeders, occur at the

March (22-26),

to the

beginning of April.

Growth, development, and molt. The young are born blind and helpless. They start to see at the age of 30-36 days. The lactation
period lasts 40-45 days. Development probably proceeds almost as
in the pine

marten and sable. At the beginning of July (Novikov,

are already nearly indistinguishable in size

from adults. By the middle of August, a young male weighed 1032 gm, a young female, 890 gm. Molt occurs twice, in spring and in autumn. Enemies, diseases, parasites, mortality, competitors and population dynamics. The pine marten, and perhaps the red fox may be considered enemies of the stone marten. Large diurnal and nocturnal birds of prey, wild cats and other carnivores are a danger to the young stone marten. A case was described when a pine marten
1962) the

young

worried to death a subadult stone marten (Rimenshneider, 1920)*.

Among

competitors of the stone marten are the European polecat,

pine marten, wild cat, domestic cat (in populated areas), and possibly other predatory animals

and

birds. In

Germany,

a case

was

observed when a stone marten killed a domestic cat (Kharbakh

[Auerbach?], 1929)*. Also noted was a daytime attack of a stone

marten on a European Polecat; the marten killed it (Frank, 1932). Diseases are not well described. In August 1 96 1 in "Vorskla
Forest" (Novikov, 1962), two young stone martens that were

obviously sick were seen: with underfur falling out and partial
paralysis of the hind limbs (encephalitis?).

Not

in Lit.

Cit. Sci. Ed.

903

603

Fig. 222.

Track and scheme of bounds of the white-throated marten on

soft

snow. Vic.

of Kislovodsk. 27 January 1950. Sketch by A.N. Formozov, about 2/3 nat. size.

In

Zaporozhsk

district of

Ukrainian SSR, the stone marten has,

in recent years, forced out the polecat,

widening the region of

its

distribution in the southern Ukraine (Ogul'chanskii, 1954). In the

Crimean preserve, a parasitic disease of the blood (theileriosis) has been observed among stone martens. Among helminthiases, the most pathogenic are crenosomatosis and mezocestoidosis of the
intestines (Rukhlyadev, 1948).

More

than 11 helminths are

known
prob-

from the stone marten (Greve, 1909). Longevity of the stone marten is not established, but
ably, similar to that of the pine marten.

is,

Among
604

Concerning population dynamics, there are almost no data. the three species of the genus of true martens, genus Martes, it is the stone marten which most easily adapts to changes introduced into nature by humans, in some places, living together with
them. In the 19th century in Germany,
it

was even noted

that the

904
stone marten excludes the pine marten. After the price of the fur
increased, the stone marten

was subjected

to intensive destruction

and, hence, the question of their protection

was

raised.

Field characteristics.
briefly,
it

When

encountered

in nature, usually

very

is

difficult to sort out those features

which distinguish

the stone marten

from the pine, the more so

to

since, in

some

places,

they are met with in similar habitats. Non-forested places or those

in

immediate proximity
it

human
talus

habitation

is

the best indicator

that
its

is

a stone marten.

It is

also said that finding the animal or

is

tracks

among rocks and

another, but this indicator

is

less reliable.

Tracks of the stone marten are distinguished by the thinner fur covering of the foot, with well developed naked food pads
(P.Yu.).

Practical Significance

The stone marten

of the

is

a quite valuable fur animal, but

Its

is

inferior, not

only to sable, but also to pine marten.

share in the fur market

USSR

is

not great: stone marten constitute no

Its

more than 10-

procurement is only in the Caucasus, in the montane part of Crimea, in the republics of Middle Asia and, in very small numbers, in the Ukraine. Due to the absence of more valuable colored fur in these places, it has quite essential significance in the budget of native market hunters. The experiment of introducing this animal into hunting allotments of Ryazan district (Lavrov, 1946) was not well thought out, and therefore unsuccessful. It was shown that the marten settled in
of that of processed pine marten.
villages there, living in bell towers and other buildings (Manteifel',

12%

1947).

The stone marten

live capture,
traps.

is

captured with jaw traps, box type traps for

sorts of [kulemka]

and various

and plate [plashka]

Hunting by shooting is not efficient. Trailing with dogs, or tracking may be successful only when the animal lies up in a tree hollow. Attempts to drive the marten away from rock slides and fissured rocks is a hopeless matter. In western Europe, it is sometimes successfully shot on moonlit nights on roofs of houses in
villages (P.Yu.).

905

Subgenus of Himalayan Martens, or Kharza

Subgenus Charronia Gray,
1865"^^

KHARZA^^
Martes {Charronia) flavigula Boddaert, 1785
1785. Mustela flavigula. Boddaert. Elench. Anim.
81.
p. 88.

Nepal.
1,

\^\\. Mustela aterrima. Pallas. Zoographia Roso-asiatica,

p.

Between the Ud and Amur

1,

rivers.

1862. Mustela (Martes) flavigula var. borealis. Radde. Reisen Slid.

Ost. Sibirien,
p.

19, 24.

Bureinsk mountains (V.H.).

60S

Diagnosis

Dimensions large larger than all other species of genus living in Old World. Tail length considerably more than half of body length. Color bright, consisting of unique combination of areas and fields of black, white, golden-yellow and brown. Sharply outlined throat patch absent. Inner part of the upper molar only slightly larger
than the outer in longitudinal diameter (V.H.).

Description
In
its

general appearance, the kharza

is

is

a true marten.

Its

typical

martenness

even strengthened and emphasized particularly sharply. It is a large robust animal, muscular and flexible, with a very elongated trunk, small, pointed head on a long neck and with
a long
tail. Its

length, together with the hairy tip, constitutes about

length.
all
is.

2/3 the

body

The

tail

is

covered with short hairs, much

thinner than in

other martens, and therefore seems even longer

than

it

actually

The limbs

are strong and relatively short with

'The proposed change of this

did not find wide application
actual requirements

name to Lamprogale Ognev, 1928 (see page 749) among systematists because it was not called forth from

and did not correspond with nomenclatorial rules. sometimes called yellow-cheastod or yellow-throated marten. This is an entirely bookish artificial name and the English translation is also an artificial name. In our language this name is also not preferred, because it is very similar to the forest marten the yellow-throated and thus leads to confusion.
^The kharza
is

906
broad feet
a long.

in

moving, the animal strongly bends

in

its

back into

hump and moves

The kharza
in

bounds which,

at a

rapid pace, are very

winter pelage has relatively short fur, which

does not have the fluffiness of the pine marten and sable and even
of the stone marten, but
it

appears lean and very long

ears are large

so than the stone marten.

are

The

even more broad, but and

short,

somewhat rounded

at their tips.

The

soles of the feet are cov-

ered with hard elastic hairs; however, the digital and foot pads are

completely naked and,

short

in general, the

paws

are

weakly furred. In

summer

fur, the

kharza appears

still

longer and thinner, with

a long, thin

tail.

Winter pelage
that
it is

differs sharply

from

that of our other

is

martens

in

relatively short, and, moreover,

harsh and lustrous, and

606 not as dense, fluffy

and compact. On the tail the hairs are short and of equal length over the whole tail. Summer fur is shorter, sparser, less compact, and not so lustrous. The color of the kharza is entirely unique. It is bright and variegated, and is sharply differentiated from the color of the other

Fig. 223.

preserve, southern Primorye. 18

Young female kharza Martes (Charronia) flavigula Bodd. "Kedrovaya" Pad December 1964. Photograph by A.G. Pankrat'ev.

907
species of the genus. The top and occiput of the head is blackishbrown with shiny brown highlights. The color of the cheeks is somewhat more reddish, which with a mixture of white hair tips produces a light gray. The posterior sides of the ears are black,
their inner portions

covered with yellowish-gray hairs as

at the

is

the hair

on the ear margin. Starting

of the body, the fur
is

occiput along the dorsal surface

a shiny brownish-yellow color with a golden

becomes browner and the sacrum and thighs are dark-brown and blackish-brown. The sides and belly have a bright yellow tone. The chest and the lower part of the throat are
tone. Posteriorly, this color gradually

rear portion of the back, and the region of the

still

brighter than the back and belly

here,

the yellow tone be-

comes orange-golden. The chin and lower lips are pure white in color. The hand and the lower part of the forelimbs are pure black, and the upper part of the extremities have the same color as the anterior part of the back. The transition from black color gradual. The tail has a shiny pure black color, except for the hair at the end
have a light violet wash. Basally the claws are white.
Individual variation in the fur
tail

hair

is

grayish-brown, the

is

considerable. This concerns

both intensity of general color and the color of the separate parts
of the body as well as the relative development (magnitude) of the
separate areas. Color of the

darker

the golden tones

The black tones
is

summer
is

fur

is

somewhat

duller and

along the back are more weakly devel-

oped. The fur of young martens

adults.

somewhat

lighter than that of

are less pure, with a brownish mixture

is

and the

throat patch

not so bright. There

no sexual differences

in color.

The

skull of the kharza in general appearance,

is

and

in a series

more similar to the skull of the stone marten, differing from it, most notably, in its large dimensions (see below). The width between the mastoid brain case is relatively shorter processes is equal to, or is a little greater than the distance from
of details,

the lower edge of the occipital

foramen
is

to the posterior

edge of the

sphenopalatine notch (ratio

region
is

is

reversed in stone marten).

The

nasal

broader and shorter. There

no isthmus

in the

middle
is

length of the nasal bones.

skull
to

The

[postorbital] constriction of the

is

behind the supraorbital process

so, than the

relatively

weak and
It

equal

the width of the interorbital space or larger.

is

larger,

sometimes significantly
canines.

width of the skull above the

The auditory bullae

are short

and widely separated

the

908
distance between them constitutes about 2/3 the length of the bulla.

The upper

carnassial tooth

is

relatively large, the [upper]

molar
is

relatively small: the length of the former is greater than the trans-

verse diameter of the

latter.

The inner blade of

the upper molar

is

only slightly broadened

its

longitudinal dimension

less than

half the transverse diameter of the tooth.

The outer
the

lateral surface

of the upper molar has a vertical groove.

male skull; age variation is significant, and generally corresponds to that of our other martens. The caudal skeleton is composed of 21-24 ver-

The female

skull

is

somewhat smaller than

tebrae (variation apparently,

is

large).

The OS penis has

is

a characteristic

form and

is

sharply distin-

guished from that of our other martens. The basal half of the bone

compressed from the sides and is quite tall, and is is noticeable on the ventral side. The anterior part rises up; its end is sharply curved dorsally so that the end is directed, not forward but vertically upwards and at the extreme tip, even a little backwards. In this way, together with the extended slightly S-shape bend of the bone, it assumes a hook form. At the extreme tip, there are four small blunt processes located in the form of a corolla, each at a corner of a quadrant. These processes are directed upwards, one of them usually being
laterally

flattened below; a fissure

larger than the others (Pocock, 1941

Indian

kharza).

In all dimensions, the kharza is considerably larger than our

608

Body length of males (9) is 500-M612-719 mm, of 500-M575-620 mm; tail length of males (6) 370M407-442 mm, of females (8) 354-M397. 0-427 mm; length of hind foot of males (7) 100-Ml 16.5-136 mm, of females (9) 100Ml 12.0-123 mm; ear height of males (4) 27-M34.5-40 mm, of females (6) 31-M37-45 mm (specimen from Ussuri Territory;
other martens.

females (12)

is

Bromlei, 1956, with additions).

Condylobasal length of the male skull

(8)

104-M109-112.5

mm,

zygomatic width of males (5) 54.4-M60.7-67.9 mm, of females (5) 53.7-M57.2-59.4 mm; skull height of males (7) 38. 0-M42. 8-47.1 mm, of females (Ognev, 1931; Bromlei, 1956; V.G. (7) 37.0-M39.9-46.8

of females (7) 96. 2-M 100.6-109.4

mm;

mm

Heptner). Length of os penis about 76-78

mm.

Weight of males
is

(eight specimens

between October and January)

(Bromlei, 1956) (V.H).

2463-M33 17-5748 gm,

of females (10 specimens between August

and February) 1155-M2765-3827

gm

909

607

Fig. 224. Skull of kharza,

Maries (Charronia) flavigula aterrima

Pall.


910
Systematic Position

The kharza, in all respects, represents a sharply isolated species even more sharply than the stone marten. Among Holarctic forms,
according to craniological features it is, apparently, nearest to the latter. Its skull is essentially a dimensional model of the stone

marten skull. Based on these characteristics its isolation from the remaining Holarctic martens is either greater or slightly greater than the stone. It is more sharply differentiated in its unique color

and OS penis structure. The latter character is considered by several authors as the most evident and important for species (Pocock, 1918, 1941). At the same time, it is the case that, given all of the
features of similarity mentioned between
all

marten species,

it

is

evidently not sufficient to separate the kharza into a separate

genus. Because of the

sum

of characters one

may

consider

it

sepa-

rable into a separate subgenus.

The kharza must be considered one of the most ancient, Pliocene forms of marten. This is supported by its peculiar range (see below). It is indicated also by the characteristic "tropical" color of
kharza which
is

absolutely not

common

to all

marten (V.H.).

Geographic Distribution
In forested regions of the

Malayan archipelago, Indochina, the

Himalayas, southern extremities of Hindustan [India], southern China and the Far East.

Geographic Range

in the Soviet

Union

This represents the northern border of the species range, and occupies the Ussuri Territory and the region of the middle, and in
part the lower

Amur. This

is

an insignificant part of the species

range.
In the west, the northern border of the range begins (coming from northeastern China former Manchuria) on the Amur, in the region where the Ol'doi flows into it (about 5330' [N. lat.] southwest of the Skovorodino railway station). At first, without noticeably separating from the Amur, orients to the middle course of the Zeya, then to the middle course of the Bureya, thence to the upper Bidzhan and Bira, including on the south in this manner, the


911

Burein mountains. From here, the border goes on to the middle courses of the left tributaries of the Amur the Kur and Urmya

and approaches the

Amur

at the

mouth of

the Khungarya.

Along

the right side of the

Amur,

the border passes eastward along

Khungarya and, crossing the mountains, extends to the ocean in the region of the rivers Koppa, Botcha and Tumnin, crossing all of
the latter.

To

the south of this described line, the kharza

is

found

everywhere, noticeably increasing in numbers towards Evidence of the occurrence of this species between the Udoi
the south'*^

and the

Amur

(Pallas, 1811)* are quite indefinite

and may fully

apply only to those places near the Amur. Information on the distribution of kharza in "Amur district north of the south slopes of
609

Yablonovyi range" (probably Stanovyi? V.H.; Ognev, 1931) and even to the upper Vitim (Kashchenko, 1913), are extremely doubtful and unconfirmed, and are not taken into consideration here. At the present time, as well as in the past, even along the middle Amur the kharza is very rare (Shrenk, 1859 and Maak, 1859, do not mention it at all). It was not recorded on the Vitim in several works of the last century (Maak, 1859). It is even absent
the

along the middle and upper Ol'doya and along the Gilyuya (Gassovskii, 1927), i.e. south of the Stanovyi range. According to
the general geographical and ecological conditions, the penetration

of this species far to the north

capture of kharza in the
the
is

is

improbable. Reference to the

Tuva

ASSR
is

("Uryankhai Territory") on

Kemchik

[.] (Ognev, 1931),

It is

completely unintelligeable and

obviously mistaken.

also entirely absent in the

Mongolian

Republic. The source of these mistakes, as in the majority of similar cases, is

probably imported skins.

Geographic Range outside the Soviet Union.

This occupies the eastern and northern parts of northeastern China

(former Manchuria), excluding, apparently, the Great Khingan, or at least, its southern part; eastern China, including Gansu and

Shensi in the west, south to Sichuan and Yunnan, with their western montane regions (eastern and southeastern edge of Tibet;
the western border of the range in

China

is

poorly known),

^'Range according to Emel'yanov, 1927; Ognev, 1931; Bromlei, 1953 and mainly according to original materials of Yu.A. Salmin and V.D. Shamykin. Erroneously cited as 1911 in Russian original Sci. Ed.


912
Indochina with Malacca, the islands of Hainan, Taiwan, Sumatra, Banks, Java and Kalimantan (Borneo). It also occupies upper Burma

and the associated Himalayan montane regions of northern India Assam, Bhutan, Sikkim, and Nepal westward to include Kashmir, Champa, Hazara (the region northeast of Peshawar) and the Peshawar region and somewhat south of it. A separate isolated part of the range is located on the west of the southern extremity of the
Hindustan [Indian] peninsula Absent in Ceylon.
(S.

Kurg, Nilgiri Hills, Travancore).

The range of

the kharza

is

very typical of an eastern Asiatic

Tertiary range. Apparently, the

this species lies in the south.

main region

for the

development of

There is a remarkably long narrow extension of the range towards the west along the Himalayas (see range of the white-chested [Himalayan black] bear). The separated section of the range in the extreme south of India, still considered,
not

long ago,

"unexplained" (Pocock,

1941)

is

typical

for

"Pleistocene ruptures" of ranges in southeastern Asia, manifested

in past periods (V.H.).

610

Geographic Variation

Up

to the present time,

many

subspecies of kharza have been

are groundless, and

described.

The majority of them

were described

609

Fig. 225.

Boundary of the distribution of the kharza, Martes (Charronid) flavigula Bodd. in the USSR. V.G. Heptner.

913

on the basis of individual deviations of color or of seasonally variable characters of fur color (it was shown that summer individuals are darker).

The form
ently, differs

existing within the boundaries of the

USSR

apparauthors

from the nominal (Nepal),

although many

express great doubt about this (Jacobi, 1922; G. Allen, 1938). There is only one subspecies in our country: Amur kharza. M.
(Ch.)
f.

aterrima Pallas, 1811 (syn. borealis).

610 Fig.

226.

Species

range

of the

kharza,

Martes {Charronia) flavigula Bodd.

V.G. Heptner.

914
It

is

distinguished from the nominal form by

its

denser and

longer winter fur and somewhat larger general dimensions. For description and dimensions, see above.
In

Amur and

Ussuri territories.

Outside the
ria).

USSR in

northeastern China (former

to the

The border with respect

nominal form

is

Manchunot known.
is

611

Differences between our kharza and the nominal form are not
sharp, but apparently are real.

lows: male (9) average

of males

431

571 mm, female 494 mm; length somewhat mm, females 408 mm. Weight
(9)
tail

Body

length of the latter

as fol-

is

less,

apparently, and also skull (Pocock, 1941).

A
melli,

series

forms described

in the area lying

between the Hima-

layas and the

Amur

(kuatunensis, szetchuensis, yuenshanensis,

similarity to the animal of the

koreand) are completely based on information stating its Amur form (Jacobi, 1922; G. Allen,

1938).

1)

Outside our country, the following forms are usually accepted, M. (Ch.) f. flavigula L. 1875 Himalayas, from Kashmir east-

f.

wards, southern China northward to Shensi and Gansu; 2) M. (Ch.) southern India (Nilgiri Hills, Kurg, gwatkinsi Horsefield, 1851

Travancore); 3) M. (Ch.)
4)

M. (Ch.)

f.

peninsularis Bonhote, 1901

and Malacca;
Kloss, 1931

5)

Taiwan; southern Tenasserim M. (Ch.)f. indochinensis Kloss, 1916 Indochinese

f.

chrisospila Swinhoe, 1866

Peninsula, northern Tenasserim; 6) M. (Ch.)

Kalimantan
all

f.

saba Chasen

et

(Borneo).

Among

the listed forms, the better distinguished ones are

the nominal, and gwatkinsii,

which

is

usually considered to be

ranked as a separate species (V.H.).

Biology
Population. Data are inadequate. Abundant on the western slope
of the Sikhote-Alin', and south to the Khor and Kkhutsinka rivers.

On

the eastern slope,

it

has already become rare in the Samarga

to
is

river basin (Yu.A.

Salmin and V.D. Shamykin). Northward

its

the extreme limits of

distribution

it

is

rare everywhere.

It

915
particularly

numerous

in the

southern part of the main axis of the

Sikhote-Alin' range.
In the southern part of the range in the USSR, the index of density of 3,2 per 1000 hectares (in "Kedrovaya Pad' " preserve,

1934-35; Korkeshko and Mirolyubov, 1936), is, apparently, not extreme; for this species, it corresponds to an estimation of "abundant". Figures for fur production are not indicative [of population],

species

is

not exploited due to difficulty and unprofitability of

hunting.
Habitat.
taiga,

Most commonly encountered

in

montane coniferous
with thickets of

although in various habitats

on rocky

cliffs

Mongolian oak,

in the broad-leaf forest zone, in

burnt-over areas,

montane spruce and spruce-fir taiga up to timberline. Most frequently, it lives throughout mossy dark coniferous forests of the Okhotsk type, along the northern slopes of the
in nut pine stands, in

and streams met with almost everywhere, appearing in great numbers in the flood plains of rivers and creeks, where it stays until late autumn. After the end of rut and spawning of the migratory fish, it becomes concentrated in montane taiga where at that time musk deer gather (Yu.A. Salmin and V.D. Shamykin). Abroad, it occurs even in the lowland swamps of Burma and
hills

in

musk deer habitat

and

in the valleys of rivers

is

(Bromlei, 1956). In summer,

it

in the arid,

unwooded mountains of the Northwestern

is

Frontier region

of Pakistan (Pocock, 1941).

Food. Kharza

an omnivorous animal, however, in the Ussuri

its

Territory the foundation of

diet

is

musk

deer, especially in the

winter time. The numbers of kharza depend on their abundance.

The kharza is a particular danger to the musk deer in winter time, when it is not possible for them to take the young of other hoofed animals, these are available to them only up to a weight of 10-12 kg. In autumn, it is no less dangerous for the musk deer when groups of nearly grown but not yet dispersed litters hunt musk
deer. In winter, the kharza tries to drive
612

musk deer onto

the ice. In

1936 on the

Armu and Nantsa

rivers,

over a distance of 200 km,

26 musk deer killed by kharzas were found (1 per 4 per 30 km. Two to three kharzas can eat a musk deer in 2-3 days (Bromlei, 1956), Food of kharza and its seasonal characteristics are shown in Table 61 (Yu.A, Salmin and V,D. Shamykin).
the carcasses of 7.7 km); in 1952, along the Sitsa river

916

Fig. 227.

Montane mixed broad-leaf

forests in

"Kedrovaya Pad'

"

Primor'e)

habitat of kharza, and also Amur badger, spotted deer, leopard and
June 1958. Photograph by A.G. Pankrat'ev.

preserve (southern
others.

Table 61. Seasonal characteristics of diet of kharza

(%

occurrence)

Type of food

Entire year

Winter
63.1

Spring

Summer Autumn
31.5

Musk and
goral,

roe deer,
calves,

45.5

44.9

42.3

moose

wapiti, wild pig (suckling)

Small exploited species

(squirrel, hare, sable,

27.1

29.7

26.6

20.0

32.2

Siberian weasel)

Mouse-like rodents, pikas

5.2

6.1

11.6

3.4
6.8
1.7

Hazelhen Small birds

Fish (migratory salmon
)

12.6

7.2

18.3
4.1

17.6
7.7

3.4
2.1

Molluscs
Insects

1.4

0.5
fruits

Nuts and

2.2

8.5

5.9
1.9

3.8

5.1

917
Ungulates and small exploited animals are the main food (20% and more) of kharza throughout the whole year, and hazelhen as well in spring and summer.
613

Small birds, mouse-like rodents, pikas, molluscs, nuts, fruits and insects rank as secondary foods in the summer period. In autumn, fish, fruits and nuts are secondary foods of characteristic
occurrence.

young wapiti, spotted Manchurian hare, white hare and flying squirrel. Mice and chipmunks are rare. Among birds, most frequent are hazelhen and pheasant. Fish, insects, molluscs and plant food occur, but are rare. In 17 data sets on diet of kharza, musk deer was found in 64% of cases, squirrel in 18%, fish in 18%, birds in 11% and hazelhen in 5.5% (Bromlei, 1956). Carrion (except fish) is not taken by kharza and it rarely returns to its
In the food of kharza are encountered:

deer, roe deer, goral, squirrel,

prey. Captures of sable are a frequent occurrence, increasing with

increase in [sable] numbers.

It

does not attack domestic fowl, but

fat

the old toothless individuals sometimes feed on discards.

Judging from the fact that kharza always possesses

its,

depos-

indicates that

it is

always well supplied with food and does not

have a "bottleneck" period during the year. Home range. The hom.e range of the kharza

is not permanent and is very extensive. In one day and night, it covers up to 10-20 km, moving in regular bounds, and in this way it can frequently travel around an entire river or creek basin. Usually, each 3 to 4 km, along the trail of a kharza, remains of eaten prey may be

found (Bromlei, 1956).

Burrows and

shelters. Information

is is

lacking.

USSR.

no information for the by day (Pocock, 1941). It travels in pairs, but in winter, litters not yet dispersed of up to 5-7 individuals are typical. The kharza hunts hoofed animals (musk deer,
In India

Daily activity and behavior. There

the kharza hunts

roe deer) in groups surrounding the prey, as observed in India. In

this

way, the yield of the hunt

is

is

increased, and the hunting of this

in

animal

determined by whether they are

it

pairs or a group. a

When

hunting ungulates,

often tries to drive

them onto

smooth

ice surface

(Yu.A. Salmin and V.D. Shamykin). In March, with weakly

crusted snow, the kharza overtakes a

since the weight load of the
to

80 gm, and of kharza

only

musk deer after 800-1000 m, musk deer on 1 cm'^ of surface is equal

31

gm

(Bromlei, 1956).

918

613

Fig. 228. Habitat of kharza in

"Kedrovaya Pad'

"

preserve (southern Primor'e).

Mixed

coniferous-broad-leaf forest with lianas. Photograph by A.G. Pankrat'ev.

614 trees proficiently. In

The kharza hunts mainly on the ground surface, but climbs moving from tree to tree, it is capable of making jumps up to 8-9 m in length. In case of necessity, it jumps into the snow from the tops of the highest trees. In captivity, it is
It is active both day and night (Bromlei, 1956). Seasonal migrations and transgressions. After March snow-

easily tamed.

falls,

the

when snow depth reaches 50-60 cm, the kharza sinks into snow and prefers to move about in the tree tops, gradually
in places
little

descending to the montane foothills to broad-leaf forests

with

snow.

Reproduction. Data are scarce. There are indications that the

kharza

is

monogamous (Yu.A. Salmin and V.D. Shamykin).

919
Increase in activity, accompanied by signs of nuptial urges,
is

ob-

served twice yearly, from the middle of February to the second decade of March, and from the end of June to the first half of

August. According to other data, copulation occurs from the first days of June to the middle of July (Bromlei, 1956). At that time
fights are

observed among males, and the animals become unwary.

Lactating females were obtained on the 4th and 7th of

contain two-three young rarely four.

May.

Litters

Young have

not been ob-

served in

summer (Bromlei,

1956).

Growth, development, and molt. There is almost no information. Summer pelage of the kharza is acquired only in August,
after a

delayed spring molt.

Autumn molt

is

usually not completed

even by the end of September. Enemies, diseases, parasites, mortality, competitors and population dynamics. The bold, strong and very active kharza has no enemies. Competitors are mainly those predators which feed on

musk

deer.

Helminth infection

is

not large.

It

lives to a consider-

able age.

Fluctuation in numbers of kharza are closely connected with

musk

The kharza population began to increase only in 1938, musk deer trapping, which caused growth in the numbers of this ungulate. However, the population increase of kharza caused, in its turn, a decrease in numbers of musk deer. Therefore, after 1941, decrease in numbers of the musk deer, were paralleled by decrease in numbers of kharza, which, in 1946, again reached the 1938-1939 level (Yu.A. Salmin and V.D. Shamykin). Field characteristics. The kharza is distinguished from other small carnivores of the Ussuri montane taiga by the large size of its tracks and its movement in large bounds (P.Yu.).
deer.
after the prohibition of

Practical Significance

The value of kharza

as a furbearer is not great

and does not comit.

pensate for the effort expended in hunting and capturing

In the

hunting industry, the kharza causes significant harm

a valuable furbearer

by destroying
on the diet

sable. In the light of recent data

of kharza, this
sirable that

harm is, possibly, somewhat exaggerated. It is dethe number of kharza be limited, but by no means
is

achieving complete destruction of this animal, which

rare in the

USSR.

920

The kharza is not easily trapped by snares or baited traps. The only method of capture is hunting with especially trained dogs. From 1936-1946, prepared skins in Sikhote-Alin' ranged from 42110 (P.Yu.).

Genus of Wolverines

Genus Gulo

Storr, 1780

1775. Gulo. Frisch Natur.-Syst. d. vierfiiss, Thiere, p.

International

Frisch' s

and the

17. The Commission on Nomenclature, has decided that book does not satisfy nomenclatoral requirements names proposed in it are invalid.

1780. Gulo. Storr. Prodr. Meth.

Mamm.,

p.

34, Tab. A. Mustela

gulo Linnaeus, 1758.

615

1780. Gulo. Pallas. Spicilegia zoologica, 14,

Pallas

p.

25 Gulo sibiricus

= Ursus gulo Linnaeus

large.

(V.H.).

Dimensions

Skull large and massive, broad and relatively short, with well-

defined rough protuberances, crests, etc. Arrow-form (sagittal) crest

high,
its

posterior end extending sharply

backward above

flat sur-

face of occiput; occipital crest well-developed, but relatively weak.

Braincase of relatively small volume. Facial part of skull short and

broad (distance between margin of alveoli of middle incisors and

middle of line uniting ends of supraorbital processes constitutes 65-70% of distance from this line to posterior end of sagittal crest).
Nasal bones short and broad, nasal foramen large and slanting
obliquely backwards in a

way

that

makes

it

strongly opened up-

wards. Orbits relatively small. Zygomatic arches powerful and very

massive, especially in posterior part, and posterior parts more widely
separated. Region of interorbital constriction quite narrow
less than
(its

width

width of muzzle above canines), elongated,

lateral sides

almost parallel. Supraorbital processes poorly developed.

Upper

profile of skull convex, considerably elevated in frontal

region and thence quite abruptly depressed

down

to nasal region.

Braincase quite narrow, but high. Infraorbital foramina small

their

diameter two times less than base of upper canine. Bony

in transverse direction.

auditory bullae relatively small, flattened, thick-walled and wid-

ened

They

are slanted as regard to

one

another, short, with swollen inner sides;

becoming wider and more

921
flattened towards the auditory meatus. Auditory canal well-devel-

oped. Hook-shaped processes of pterygoid bones not fused with

auditory bullae. Mastoid (mammilary) processes large and protude

obliquely forward anteriorly and

downwards beneath

the auditory

meatus; paroccipital

(lateral

occipital) processes large, separated

from bony auditory bullae. Bony palate very broad. Dental formula as in true martens, genus Martes:
3 14 = 38. I-C-P-M1

14

First

premolars sometimes absent, with socket of tooth not

evident. In

some

cases, all four [first premolar] teeth absent. Teeth

large and powerful, relatively

much

stronger than those of true

martens (genus Martes). Longitudinal diameter of base of upper

canine about 10

mm,

usually larger (canines significantly

more
in

powerful than those of badger).

Upper molar

typical marten-like in structure

its

elongated
(its

transverse direction, with

inner blade a

little

wider than outer,

longi-

but small. Upper carnassial tooth very large and strong

tudinal diameter exceeds that of the upper molar,
times), but the inner cusp in
its

anterior part

is

by nearly four relatively weak.

lateral

The main apex of this tooth is high and massive, with sharp cutting edges. Lower carnassial tooth large, with two massive
of almost equal height.
First premolars of both

apices

616

jaws very small and somewhat crowded and second lower premolar also very small, with a rounded crown. Upon closure of jaws, crowns of second and third upper and of third and fourth lower premolars do not overlap they are located at different levels, or they only touch each other or barely come together (third and fourth)'. The longitudinal axis of the upper carnassial tooth is parallel or almost parallel to the in all other genera represented in longitudinal axis of the skull
into toothrow,

our country, these lines form an angle,

in

some

quite large.

Os penis

quite massive, weakly curved in the middle part,

gradually thickening in basal half and thick at base. At anterior

thin end, there is an expansion in
anteriorly.

form of a

fist,

slightly divided

'Indications are encountered in literature that in the closed jaws the apices of the

carnassial teeth (upper and lower) "are widely separated" (Ognev, 1935) were based

on misunderstanding

they

form

tightly closed scissors.

922

Trunk short and massive, limbs of moderate length with large

broad
feet, especially in forelimbs,

semiplantigrade; claws large.

relatively small, ears

Facial part of head

somewhat extended, eyes

small, short and rounded. Tail short, without terminal hairs,

approximately equal to length of head. Winter fur very long, dense and shaggy, with brown tones. Seasonal dimorphism in fur character sharp, in color weak; sexual dimorphism not observed. Sexual differences in measurements quite considerable. In addition to paired anal scent gland, there are special glandular regions on the belly
in front of the sexual opening,

developed

in

both sexes^.

Two

pairs

of nipples.

The range of

the genus

is

very large, occupying the taiga and

tundra zones of both the Old and

New

Worlds. In the

past, the

range extended considerably farther to the south (for details, see

below under species description). Genus Gulo constitutes, within the family, one of the most peculiarly characterized genera. Its independence has never been
doubted.
It is

a sharply characterized genus, not only morphologi-

cally, but also ecologically

and zoogeographically. However, the

separation of the wolverine into a separate subfamily cannot be

considered well-founded. Such a point of view one would be able

by admitting into consideration fine details of the entire marten family (more than 10 subfamilies), a fact (see above, family characteristics) which does not seem necessary in the systo hold only

tematic relationships of the family as they are described

present time.

at

the

Among

the real features of the genus,

it

is

not posall

sible to distinguish those which,

against the background of

genera, could have supergeneric significance.

to to

Concerning systematic position, the genus is definitely closer the true martens of the genus Martes. They are closely related
each other not only
in dental

formula and a series of features of

skull structure, but also in such characteristics as the unique glan-

dular area on the belly and a series of

homologous

instincts

and

behavioral features.

On

the whole, in

all

of their characteristics,

martens stand closer to the wolverine (Krott, 1959) than to weasels

and

polecats, with

which

it

is

usually placed in close proximity.

in the general

Only some, actually secondary features

appearance

^The secretion of the anal scent gland of wolverine can be smelled for a distance
of 3 m. The glandular field on the belly serves to mark the territory. of the family,
it

Of

all

species

is

characteristic only for marten of genus Martes,

and for wolverine.


923
of wolverine give an impression of great uniqueness of this form
in actually a gigantic

marten (Pocock). Attempts to

affiliate the

wolverine with the South American Tayra (Eyra) barbara (Krott,

1959) are lacking in serious foundation.

The genus Gulo

is

also related to the genus Martes in origin.

The genus is known from the lower Pleistocene of Eurasia and America (G. schlosseri). Its ancestor was, probably, the closely related Pliocene Plesiogulo (Perunium), known from the lower
Pliocene of Eurasia (P. monspessulanum, P. brachygnatus) and
the

middle Pliocene

in

America (Simpson, 1945; Thenius and

Hofer, 1960). Separation of the lineage which gave rise to contem-

porary Gulo from the genus Martes, which arose in the Pliocene
(perhaps in the upper Miocene) occurred, evidently, in the Miocene.

The genus Gulo

itself

appeared in the lower Pleistocene.

Tertiary wolverines were thermophilic, and only during the

course of the Pleistocene era did the range of this group receive
recent shape
618

its

taiga

and forest-tundra, and

in part,

even tundra.
as

Quaternary wolverines have been derived from the lower Pleistocene

G. schlosseri and they

do not

differ

from recent wolverines

regards species relationships (see below concerning distribution of

wolverine outside the USSR).

In the

genus there

is

only one species: G. gulo (Linnaeus,

1758).*
Specific identity of animals of the Old and

New Worlds

is

evident, although several authors at the present time usually consider the wolverine of North

America

as an

independent species

(G. luscus L.^).

In the

USSR,

there

is

one species

the

wolverine G. gulo

(Linnaeus, 1758)*.

The range occupies

tundra zones.

the taiga

and a considerable part of the

As

a fur-bearing animal, of secondary importance, but is in

part injurious to the hunting

economy

(V.H.).

Parentheses omitted
ingly

in

Russian original

Sci. Ed.

'The placement of the Holarctic wolverines in one species was recently convinc-

shown by special investigation carried out on reliable material (Kurten and Rausch, 1959).

925

WOLVERINE [ROSOMAKHA]^
Gulo gulo (Linnaeus, 1758)*
1758. Mustela gulo. Linnaeus. Syst. Nat., ed. X,
1, p.

45. Northern

Scandinavia (Lapland).
1780. (Ursus) luscus. Linnaeus. Syst. Nat., ed. X.
1, p.

47. North

America, Hudson Bay. 1780. Gulo sibiricus. Pallas. Spicil. Zool.

Ob'

river.

Requires confirmation

14, p. 35, tab. 2.

Upper

Altai,

1792. Ursus gulo albus. Kerr. Anim. Kingd. Syst. Cat. No. 381,
190. Kamchatka.

1816. Gulo vulgaris. Oken. Lehrb. Naturg.

ing of Gulo.

3, 2, p.

1004.

Renam1, p.

1820. Gulo borealis. Nilsson. Skand. Fauna. Dagg. Djur.

95.

Renaming of Gulo.
1829. Gulo arcticus. Desmarest. Mammalogie, 174.

Gulo.
1829. Gulo arctos.
1,

p. 68.

Renaming of Gulo.
Mountains south of Telets Lake,
Altai.

Renaming of

Entw. Gesch. Nat. Syst. Europ. Thierw.

1918. Gulo biedermanni. Matschie. Sitz-Ber. Ges. naturf. Freunde.

Berlin, p. 147.

1918. Gulo wachei. Matschie. Ibidem,

river north of Belukha, Altai.

p. 147. Sources of Katun'

1922. Gulo kamtschaticus. Dybowski. Arch. Tow. Nauk.

p.

Lwow,

1,

349.

Nomen nudum.

1948. Gulo gulo kamtschaticus. Averin. Trudy Kronotsk. gos.

zapovednika, 1, p. 145. Nee Dybowski, 1922. Mouth of Povorotnaya river, Kamchatka.

Sometimes the name of

all to

this

animal [rosomakha] actually somewhat unusual,

is

written with a double "S" [in Russian].

At the same time,

this

word has nothing

at

do with the root "ross". According to Dal', this is the true name of the animal and it should be written as given above. It is also sometimes used as a word of abuse, meaning "scatter-brained, sloven", in those regions where the wolverine does not live and the people do not know it (Kursk, Ryazan, Tambovsk, former Simbirsk districts). The expression "walks like a wolverine" means to walk "wearing his clothes unbuttoned" (Dal'). The habits of this animal allows us to assume that its name has been derived from the Russian common language. In the languages of the people of Scandinavia and the northern USSR, corresponding root is absent.

926

Diagnosis

The only species of

the genus.

Description
In general appearance, the wolverine is very distinctive, not similar to

other

members of

the family, and

is

particularly strongly

distinguished from the true martens and the species which are
externally similar to them.
619

In winter fur, the wolverine is an animal of heavy,

massive
approxi-

appearance, short and broad, especially the posterior part of the

trunk, on strong short legs

and very broad

It is

feet. Tail short,

mately equal to length of head.

fur. In its

clothed in long shaggy hairs,

weakly demarcated from the croup which is covered by dense long general appearance, it reminds one of a bear. The head, however, is relatively small, covered with short hairs and does not appear thicker than the neck. It is moderately elongated, with small widely separated ears with rounded tips. They slightly protrude from the fur. On the whole, the head in form does not have the
pointed shape characteristic of martens.

..>"

Fig. 230. Wolverine,

Gulo gulo

L. Sketch by

A.N. Komarov.

927

The wolverine

is

a strong, clever

and

alert animal, but

It

conveys
in

the impression of being heavy and clumsy. jumps, somewhat laterally and as if stooping the trunk
is

usually

moves
its

the posterior part of

all

higher than the shoulder. Although

of

move-

ments are quick, they seem clumsy and uneven, which is still more accentuated by the undulations of the long fur. In summer fur, the wolverine looks less massive but, if you like, even more clumsy. Still more striking to the eye are the thickness and strength of the legs, and the head looks much larger. In winter fur the feet are densely furred, but in summer, the naked areas on the lower surface of the feet are easily seen.
their tips;

Claws
is

are light-horn, lighter at

on the foreleg

their length

24-26 mm, on

the hind

22-24 mm.
Pelage of the wolverine
dense, length 30-35
in

winter

is

pactly arranged but "shaggy" and harsh.

dense and long, not comThe underfur is not very

and not hanging down. Guard hairs are short only on the head and paws and are quite compactly arranged. Their length on the forehead is 10-15 mm, between the ears about 52 mm, on the back 80-100 mm, and on the shoulders and
thighs

mm

120-150
seems

mm.
to

Therefore, the animal's trunk especially

posteriorly,

hanging, undulating hair, imparting to the trunk in no

its

be enveloped by a band ("skirt") of long little degree

above the ankle and wrist articulations, the hairs have a length of 80-100 mm, and on the paws 25 mm. Body hairs weakly adhere to each other and stand at a large angle to the skin surface, marking the fur seem very fluffy. Tail hairs are extremely dense and have a length of 65-280
peculiar appearance.
the legs,

On

mm.
620
it

Therefore, the

tail

is

very thick and fluffy, although short.

The fur of wolverine is of low value due to its coarseness, but one of the most is warm, sheds water well and is very durable

durable ("long- wearing") of furs, close in this respect to

otter.'

Summer

fur

is

scarcer and shorter than in winter, but the rela-

tive difference in hair length

on different parts of the body

is

retained, hair

on the

tail

being very long and dense.

'Apparently, therefore, guard hairs are very fluffy and the underfur does not hang down, wolverine fur in part possesses a special property which is absent in other furs: the hoar-frost from breathing which settle on it is easily shaken off and the fur does not wet. This is very appreciated by several northern people who make collars of winter clothes

from wolverine
arity

fur.

There,

much money

is

paid for skins of this species. This peculiin the Arctic (Krott,

of wolverine fur was confirmed by investigations of the American military de-

partment,

who

use

it

for special

uniform parts

1959).

928
Spring molt
is

complete, but in autumn probably, there

first

is

no

shedding (completion of hair growth). In the is fully covered with adult pelage.

autumn, the animal

The color of

the wolverine

is

entirely species-specific. In ani-

mals of more or less average color type, the anterior part of the muzzle is dark brown. On the forehead, occupying the whole area between eyes and ears, and sometimes even descending to the

sometimes whitish or yellowish-white field, due to the presence of light ends of guard hairs. Ears are dark-brown, slightly lighter on the inner side. The occipital region is brown, the dorsal neck dark-brown, slightly lighter on the sides. The top of the neck, withers and the whole middle part of the back to the sacrum are occupied by a uniform color field of dark brownish or dark-chocolate color. The guard hairs have shiny
zygomatic region,
is

situated a lighter,

dark-brown ends, the deep underfur

tinges.

is

brownish-gray, with ashy

is

The general tone of

this

"saddle"

very beautiful, deep

and bright. Extending from the sides of the neck, through the shoulder region along the sides of the trunk and further including the rump, are two light bands. They are at first narrow, not sharp and not bright, but passing posteriorly, they broaden and become lighter. They include also the hairs on the top of the proximal third or half
of the
tail.

This pattern ("breast-band")* has a light-tawny color

and

is

well demarcated on the general dark color of the animal.

The whole lower part of the body, beginning from the chin and below the shlei, is covered with dark-brown or blackish-brown fur,
darker than the saddle. Elongated white spots with irregular outlines often occur in the

middle of the [ventral] neck and on the

chest between the anterior limbs.

The

legs are covered with shiny

blackish-brown hairs
part,

they are darker than the saddle and the legs

The
tail,

are near-black or black.

except of the above-mentioned

has a color corresponding to that of the saddle, or a somewhat lighter reddish and dirty color. In the coloration of the
animals of
this type, the light tones are strongly

developed but

cede to those occupied by dark tones.

Wolverine coloration is subject to great variation, partaking mainly an individual character, but also, although to a lesser
*The Russian word "shleya" is the name for a sailor's harness that crosses the and over the shoulders, continuing down the sides, and is attached to rigging, enabling the sailor to haul ropes attached to the ship's rigging more efficiently Sci. Ed.
breast

929
degree, geographical (see below). This variation
is

demonstrated

both in general color tone and

in

form, size and generally, degree

of development of the breast-band,

in the

deviations from the type de-

scribed [above] are very great. Very dark wolverines (dark-brown

terminology of furriers) have a very dark dark-brown saddle

all

remaining body parts are darker than described. The breast-band begins farther back, is narrow, and does not include,
tone, and

or nearly include, the

tail

base; colored only a

little

lighter than the

saddle and shows weakly on the skin, mainly in the posterior part of the body. In animals of this type, the light tones are reduced

both in area and in color. In extreme cases of darkening, the ani-

mal looks almost evenly colored. The frontal

fined or almost unnoticeable.

field is

weakly dehave the

is

Wolverines of the
dark part of the skin
is

light color type ("light-brown")

saddle of light-brown color, and general tone of the remaining

correspondingly

light.

The breast-band

well defined, distinctly observable on the broad shoulder blade,

particularly posteriorly, and covers a considerable part of the
621
It is

tail. is

light-ocherous or even whitish.

The

frontal transverse

band

sharply defined and very light. In animals of this type, light fields
are generally

no less developed than dark (not considering

occupy
a very

abdomen).
In the extreme cases, the light parts of the skin

large area, and have sharp outlines and very light straw-yellow
color.

The dark

areas are greatly reduced in extent and color

they

on the sides of the neck, the light stripes of the breast-bands unite between the shoulder blades, forming, in this way, a broad, completely closed
are light, reddish-brown or brownish-red. Starting
ring.

Since the stripes of the breast-band are very broad, the saddle
represented only as a small rounded dark field in the middle

itself is

of the back, surrounded by a broad light area. The forehead and

bridge of the nose are whitish. In this type of wolverines, the light
tones exceed the dark both in extent and tone.
In dimensions the wolverine
is

one of the very largest species

of the family.
is

Body
tail

length of our wolverine (males and females)

70-105 cm;

length without terminal hairs, 18-23 cm; length

of hindfoot without claws, 17-19 cm; ear height 5-6 cm; and

shoulder height 35-45 cm.

1.5 years) of

Condylobasal length of the skull (old and adult, i.e. older than males is 132.7-158.0 mm, of females, 128.7-149.0

930
zygomatic width of males, 92.7-110.0 mm, of females, 89.0mm; interorbital width of males, 38.7-44.2 mm, of females, 35.0-42.3 mm; length of upper toothrow of males, 51.0-59.7 mm,

mm;

103.0

of females, 49.0-56.8
(after

mm. Weight

in

winter time

is

10.7-19 kg

Stroganov, 1962, supplemented by material from Z[oologicaI]

of]

M[useum

M[oscow] U[niversity] and other

sources; total about

100 individuals)'.

The measurements
ation

are subject to quite significant sexual varilighter than males, with this differ-

females are smaller and

is

ence being perhaps quite significant.

Some geographic

variation in

dimensions

present (see below).

Morphological characteristics of the wolverine are explained

by Krott (1959)
in the past as

in his

monograph of
is

this species,
is

by the

fact that

well as at present, the animal

is

associated with bogs.

This opinion
author's

incorrect and

explained by the fact that the

own

observations, carried out in a very limited territory in

the northern part of the Scandinavian Peninsula, were mechanically extrapolated to the entire range.

As

is

evident from the bio-

logical features of wolverine to be brought

up

in notes

below,

it

has no particular connection with bogs

at all.

Some

of the peculiar

features of the animal (foot structure, fur characteristics and

others)

show
is

a strong

and ancient connection with snow. Also

a

incorrect

the idea of the cited author, that the connection of this

itself represents
is

animal with bogs

nist.

known

sort of biological
its

vicariance with the wolf, which

In actuality, the picture
is

believed to represent

antago-

quite the opposite

carcasses

of
life

animals rent by wolves (moose, reindeer) facilitate the winter

of wolverine.

connection of wolverine with reindeer has been

it

noted thusly;

in several places

vanishes together with the disap-

pearance of reindeer (V.H.).

"Information given in the monograph of Krott (1959); weight

vidual males up to 35 kg, females not

20-25

kg, indi-

more than

2/3 weight of males, often half, are

incorrect. Information in our literature about 32 kg wolverines in weight (Novikov,

1956) are encountered but are doubtful to the highest degree. Even the autumn badger,
heavily fattened before hibernation, are rarely more than 20 kg in weight. The Ameri-

can wolverine weighs 10.8-18.2 kg (24-40 pounds; Hall and Kelson, 1959). Females
are less than males by 10 to

15%, but never by a

third or half.

931

Systematic Position
iJOSJi.

622

Fig. 231. Skull of the wolverine,

Gulo gulo

L.

932
623

Geographic Distribution
The
forest zone,

mainly

taiga, of Eurasia

and North America, and

Worlds.

in part the tundras of the

Old and

New

Geographic Range
This
is

in the Soviet

Union

very extensive and constitutes more than half the range of

all

the species and almost

of

its

Eurasian part.

It

occupies

all

of

Siberia and the northern half of the European part of the Union,
i.e.

the greatest part of the State's territory.

Establishing the precise northern limit of the range

cases, difficult since the wolverine

is,

in

many

roams very widely. Being,

strictly

speaking, a forest animal,

it

travels very far into the tundra,

sometimes for hundreds of kilometers. Cases are known when wolverine wandered 250 km around the tundra and mountain ranges for two weeks (S. Naumov and Lavrov, 1953). In the forest-tundra, it lives a more sedentary existence and reproduces everywhere, and in several places it moves into the southern part of the tundra. At the same time, information on the distribution of wolverine and on the northern limits of its range are so very rare that it is
difficult to differentiate the region of its

normal and permanent

existence and breeding from the region of transitory occupation.

The northern border of

stipulated, are data

the range given below, if not otherwise

on the most northerly occurrences, including

transgressions.
In the west, the northern border of the range passes along the

ocean coast of the Kola Peninsula (Pleske, 1886) and along the White Sea eastwards at least to the mouth of the Mezen'. At Kanin, it goes to approximately latitude 67 N. lat. (Zhitkov, 1904), but its transgression to Cape Kanin is very probable. Farther eastward,
the range border passes along the shore of the Bering Sea, but in the northern tundra regions, litters are very rare, and the region of

normal reproduction
belt

is, apparently, connected with the krummholz and perhaps with the very southern part of the tundra (Leble, 1953; V.Ya. Parovshchikov).

In the northern Ural region, the border extends to the sea

(Shvarts, Pavlinin and Danilov,

1951) and includes the Yamal,

the extreme limit of

passing along the Malygin Strait

ing approximately 73 N.
lat.

(Zhitkov, 1913), therefore reachis its

This

usual

933
transgressions; however, a den

was found

in the southern

Yamal

at

Yarro-to Lake (about 68 N.

lat.,

Zhitkov, 1913).

Eastward, the boundary also includes the Gydansk Peninsula

(S.

Naumov, 1931)

and, bypassing on the south the islands of this

part of the

Kara Sea, passes onto Taimyr. Here, the border passes

its

very far to the north of the peninsula (north of Taimyr Lake) and
nearly reaches

northern

tip,

probably including
in

it

entirely.

The

northernmost occurrences of wolverine

Taimyr

are:

lower Kheta

frequently); Omulev Bay in the Yenisei mouth of the Khur, left tributary of the Pyasina (73 N. lat.); Novaya and Bol'shaya Balakhnya rivers (about 7320' N. lat.); (Yakovlev, 1930) and Taimyr Lake at 74 N. lat. or somewhat farther north. At the mouth of the B[olshaya] Balakhnya it is, apparently, extirpated (V.N. Skalon). This is the most northerly place of breeding in the Old World. Finally, the wolverine was recorded (in migration) along the northeastern Taimyr

and Khatanga (72 N.

at

lat.

Gulf

7240' N.

lat.

coast northwards to the latitude of the Faddei islands,

lat.

i.e.

to 77

N.
the

or a bit to the south (E.I. Shereshevskii), and the northern

is

extremity of Chelyuskin Peninsula (Rutilevskii, 1939). This

northernmost occurrence of the species in the Old World.

To the east, the border passes along the ocean coast (A. Romanov, 1941) and includes the mouth of the Lena (Cape Bykov; Kolyushev, 1936) and the delta of the Yana (Bunge, 1887). The
wolverine
is

known from

the
(E.I.

New

Siberian islands

from

the

Bol'shoi Lyakhovsk Island

Siberia (Tugarinov,

Shereshevskii) and even

Novaya
islands,

Smirnov and Ivanov, 1934). On the

the wolverine occurs only as a migrant, crossing the ice of a strait

about 50

km

in width.

Information on the distribution of the wolverine farther to the

east, are exceptionally rare.

Apparently, the boundary passes along

the coast and probably reaches the

mouth of

the

events, in this region, the wolverine lives along

625

Kolyma. In Omolon, and

its

all

the

Bol'shoi and Malyi Anyui. Beyond, the range extends over the

whole basin of the Anadyr'

to its

mouth, including

northern

Chukotska (Anadyrsk) range (Portenko, 1941). The range even includes the Chukotsk Peninsula (Grinberg, 1933) and probably reaches its northern coast. The Pacific Ocean comprises the eastern border of the range.
tributaries, and, apparently, at least to the

The wolverine

inhabits Sakhalin and the Shantar Islands (at least on Bol'shoi Shantar and Medvezh'e; Dul'keit, 1927) and is absent on Karaginsk, and the Commander and Kurile Islands.

935

The outlined area of

of the Far East
tory;
is

distribution of the wolverine in the south

is

poorly revealed (information

a quite

partly contradic-

Ognev, 1935) and, apparently has

it

odd form. In Ussuri

Territory, the range occupies the Sikhote-Alin' [range], descend-

ing along
it is

approximately

to

44 N.
is

lat. (in

Sudzukhinsk preserve,

absent; G.F. Bromlei). This

the southernmost point of oc-

currence in the Old World. Thence, the border passes northward

along the western slope of the range, reaching the Amur, apparently,

somewhere
it

in the

region of the mouth of the Gorin.

From

upper parts of the Kura, and passes to the lower Bureya and Zeya (Arsen'ev, 1923; Ognev, 1935), and somewhat to the west, exits at the boundary with northeastern China (formerly Manchuria). From the Zeya to the taiga regions of the southern Altai and Markakol
here,

directs itself westward, across the

Urmya and

Bira,

Lake, the southern border of wolverine range in the

outside of the

USSR

passes

USSR

or extends along

its

southern boundary (Tannu-

Ola range; Tugarinov, 1916; Yanushevich, 1952). The animal is absent only in the steppes of eastern and western Trans-Baikal. From Markakol Lake, the border (reconstructed), surrounding the Altai from the southwest and west, passes through the Kurchum, Zyryanovsk, Ul'ba and Leninogorsk regions (Riddera; Sludskii, 1953), then bypasses the Tigeretsk range and then turns again to the east, enclosing the Altai from the north, and then proceeding westward and northward to the Kuznetsk Alatau and, apparently
also including the Salairsk chain.

From

the northern portion of this

range and the Novosibirsk region (Berger, 1946; Laptev, 1956),

the southern border of the range suddenly turns

westward and

proceeds to the Urals along the band of forest-steppe through Ghana

Omsk and Kurgan or the region between Kurgan and Troitsk (Kurtamysh southwest of Kurgan, Talovk and Shchuch'e west of it and other places here). In the Urals, the border passes through Chelyabinsk to Zlatoust and Satka (Kirikov, 1959). Along the Ural, the range extends, southwards approximately to the latitude of Zlatoust (Sabaneev, 1874) and even to Sterlitamak (about 5330' N. lat.; Eversmann, 1850). In the Urals and in the European part of the Union, the range has been very greatly changed during the last century. At the same time, old information about it is very poor and the boundary of the reconstructed range might be conveyed only approximately.
Lake,

936

As with ranges of

several other taiga

mammals

(reindeers),

it

is

probable that they extended farther southward than at present.

The range boundary ascends abruptly northward along the western slopes of the Urals, to and somewhere near the 56 parallel, it

abruptly turns again to the west, going along the

Kama

and

Volga, seemingly to around the mouth of the

passing north of
ago,
it

Oka and

farther,

probably, along the more northerly part of the Oka- Volga interfluve,

Moscow.

It is

most

likely that, a very long time

proceeded more southerly,

i.e.

included

Moscow

province

and the entire Volga-Oka interfluve, in particular the Meshchera and several adjacent places, but in surveying that time verifiable
information on the past occurrence or appearance of the wolverine,
for

example

in

Vladimir, Ryazan and

Moscow

districts, is absent.

For the Gorkii district, there is information on the occurrence of wolverine in the past only in the northern forested parts of the
former Makar'evsk and Semenovsk counties (Puzanov
i.e.

et al.,

1955),

not south of the Volga.

forests in the region

local, in its 626

may be assumed that throughout the between Moscow and the Urals, the range was
It

southward extent, for example, along the Sura.

the

West of

Moscow
It

meridian, the range descended signifidistrict,

cantly southward.

occupied Smolensk
parts,

apparently, to

its

extreme southwestern

and probably included the extreme

Byelorussia

western parts of Bryansk

district also (Klintsov region),

and

all

the Baltic republics, including Lithuania and, probably,

district.

Kaliningrad
(Brinken,

The most southern place of occurrence

west was along the right bank of the Dnepr

Belovezhsk

in the

Forest

1828) to the Pripyat' (Pinsk woodland), the Ovruch

mouth), west of Kiev

region (former Ovruchsk county), the Pripyat'-Usha interfluve

(flowing into the Pripyat' not far from
near
its

Radomyshr (border of former Kiev and Radomyshl' counties), Kanev region (former Kanev region, i.e. near the Dnepr
directly south of 50 N. lat.) and, finally, the city of Smotrich

(about 40

km

north of Kamenets-Podol'sk). This

is

the southern-

most

point, apparently, of the

more or
at the

less

permanent residence of
lat.).

the wolverine in the 7th cent, (about 49 N.

Farther south,
to

only transgressions were

known
is

end of the 18th century

Davydon Brod on
about 4715' N.

the Ingulets river (100

km

northeast of Nikolaev;

most southern point in the European part of our country where the wolverine was recorded.
lat.).

This

the

'

937
In the past, the wolverine

bank

of

the

woodlands"

was not encountered along the left of Chernigov ("Chernigovsk the northern forest parts of the former Chernigovsk

Dnepr

south

governance).

As regards the extreme western parts of the Ukraine, Carpathian and Ciscarpathian, information is absent, but it is possible that very long ago, wolverine occurred in forest areas even here. The
State boundary and the Baltic Sea

the coast

constitute the

although the range never reaches

its

western border of the range.

distribution, the wolver-

Therefore, at the southern limits of

ine

was

also found in the forest-steppe and transgressions into the

it still

steppe zone occurred (where

occurs at present
its

see below).
great,
fell-

The range of
was subjected

the wolverine, and especially

southern border,

to quite significant,
last century.

and

in
is

some places very

changes, during the

This

related to both the direct

extermination and exclusion of the animal, but chiefly to the

ing and thinning of the forests. At the present time the wolverine,

which was also previously found

coniferous forests,
is

in

regions of broad-leaf oak-dark

now

restricted in its distribution to taiga

coniferous forests. Therefore, the southern boundary of the range

European part of the Union retreated northward very for more than 100 km in the extreme west, and in some places, to considerably more than the border of the corresponding zone shifted. In the east, recession was also significant, but much weaker. Moreover, in the range, which was never more or less continuous, significant "hatchways" were formed locally, associin the

strongly

ated with changes in natural conditions.

In Siberia, changes in the southern part of the range also

occurred; however, they were fewer and there

is

very

little

material

on them. In middle and eastern Siberia, the range, as before, reaches

the State frontier, locally there are large "gaps".

The southern

border was also not subjected to substantive changes in the Far

East.
its

north, there are

Concerning the distribution of the animal in the Altai and to no data. In western Siberia, the southern border

of more or less permanent occurrence of the wolverine in the 50'

passed, apparently, along the line:

Tomsk-Kozhevnikovo-Baksa
On

river

(southwest of
map

Tomsk

upper Om' divide between the Tartas and

after S.V. Kirikov (1952, 1959, 1960).

Places of occurrence
(1960), instead of

S.V. Kirikov's

Davydov Brod (this point is not plotted), a transgression to the Azov Sea coast south of Melitopol' was indicated. This is a clear cartographical error
is

since in the text there

no reference of

this sort.

938
(data of Stroganov, 1962)

Tara-Tobolsk-Irbit.

In the Urals,
Irbitit

the southern border of the range passes along the line:

Egorshino-Perm' (Shvarts, Pavlinin, Danilov, 1951). Farther on,

627

goes a

little

north of Kirov, again ascends northward in the Nikol'sk

region (to the

Tot' ma (at

Yug

river south of Velikii Ustyug), goes farther to

60" N.

lat.)

and

at

approximately 60 N.

lat.

crosses the

whole of Vologoda district. In the recent past (about 1930), the wolverine was encountered in this district and south of the mentioned parallel (Gryazovetsk region south of Vologoda; Savinov and Lobanov, 1958). To the west, the border reaches the Gulf of Finland somewhere in Leningrad district (Novikov, 1956). There have been corresponding changes on the western border of the range; northern and eastern were not affected. The range reduction referred to took place mainly in the last century. Thus in Lithuania, Byelorussia and the northern Ukraine, the wolverine was, apparently, still a normal member of the fauna 150-200 years ago (Kirikov, 1952), and in Belovezha Forest, it was met with about 1800 (Brinken, 1828). The complete disappearance of wolverine in Byelorussia dates from the 90' s of the previous century, when the last animal was killed not far from
Slutsk (Fedyushin, 1929).

At the end of the 19th, and

vations of wolverine were

in our, century, individual obser-

known

south of the described line.

one hand, they characterize the process of range reduction the capture of the last remaining animals, as in the eastern half of the European part of the country and on the other hand, they repre-

On

sent occasional long-distance transgressions of single animals.

They

reach not only the southern border of forests, but also the foreststeppe region and even the steppe and, in exceptional cases, the
semidesert. For example, a transgression to the Kly river (north of
the western tip of Zaisan Lake, in 1949, and in the 30' s, of this

Lake Ghana (Ghanov, Kuibyshev and Vengerovsk regions of Novosibirsk district, 19481951), to Ghagly Lake north of Kokchetav (1949; Sludskii, 1953, 1953a), to Borovom southeast of Kokchetav (Mikhel', 1934), to Zlatoust on the Ural (Shvarts, Pavlinin and Danilov, 1951), in the northern parts of Tatariya and to Kazan (Grinberg, 1933)^ to
century, semidesert) in the area north of
'Lake Chagly is surrounded by steppe for 10 km. information on transgressions in the 20's into Ivanov and Moscow districts (Grinberg, 1933) are doubtful and are, apparently, based on cases of prepared imported skins.

939

Gor'kov
ties

district (1943;

Puzanov

et al.,

1955), in different locali-

in

Yaroslavl' district, in particular the

district (1925), to

Lyubim

region, in

Bezenberg in Estonia (1890), in the El'tsa region on the Sosna river (1925; Ognev, 1935), to the north (Dmitryashev region; 1940) and southwest (Verkhyaya Devitsa river in Khokhol'sk region; 1952) of Voronezh (Barabash-Nikiforov,

Novgorod

1957), to the Slutsk region in Byelorussia (see above), in the northern parts of Kiev district and to Kiev, in

Radomyshlya (1914) and

series of others.

Proskurov (1924; Sharleman', 1952)* regions and a

Geographic Range outside the Soviet Union (see map

description)
In North

in

genus

America extends

to the northern shore of the continent

(except, apparently, a narrow strip of the Arctic coast of Alaska)

and occupies the entire Arctic archipelago, including Ellesmereland north of (the northernmost point of the species range as a whole 80 N. lat.), except for Banks, Prince Patrick, Sverdrup and several

others in the northeastern part of the archipelago (Hilzheimer,1930);

the wolverine
is

absent in Greenland.

The southern border goes through

Vermont,

New

Brunswick, Maine

New

York, Pennsylvania, northern Ohio, with an exten-

sion to the south into southern Indiana and thence passes to the

northwest to northern Minnesota and North Dakota.

as far as the Pacific

From

here and
in the

Ocean, the range passes far to the south

montane regions as large extensions. One narrow extension passes from North Dakota and Montana to western Nebraska; another from Montana and Idaho as an extension of complicated outlines extending, in a divided manner, to Utah and Colorado; a third, inclining toward the coast of the Pacific Ocean, passes at first along the western parts of Washington, Oregon and northwestern California and then, as a long extension, along the Sierra Nevada almost to 35 N. lat., or a little to the south. This is the most southern point of occurrence of the species. The range includes Kodiak, Vancouver and several other Canadian islands in the Pacific Ocean. On the Atlantic coast, the range occupies Newfoundland, but Anticosti, Prince

Edward and Queen Charlotte Islands**

are not included in the range.

*Not in Lit. Cit. Sci. Ed. **These are off the Pacific, not the Atlantic, coast

'

Sci. Ed.

940
628

In

the

Old World, the reconstructed range occupies the

(in the

Scandinavian Peninsula
part),

extreme south, transgressions oc-

cur in recent times), Finland (at present, absent in the southwestern

and central Europe

to

land (in the mainland of Europe,

Braunschweig and Saxony and to Poit has been absent for a long

time). In Asia, outside the limits of

USSR,

the range extends southIn north-

wards

to the northern part of the

Mongolian Republic.

eastern China (former Manchuria), the range occupies the Great

Khingan (probably, its northern part) and Ilkhuri-Alin mountains (Baikov, 1915; Lukashkin and Zhernakov, 1943). Its occurrence in Lesser Khingan was recorded (Baikov, 1915). In the Mongolian Republic the range occupies Kentei down to Ulan Bator and even a little to the south (about 4745' N. lat.) and the PriKosogoF
mountains.
In the Pleistocene

and Holocene, the wolverine was found

outside the region of occurrence outlined above; in historical times

it

in

was encountered in Poltav and Voronezh districts, in the Crimea, the Caucasus (Trans-Caucasus), in Hungary, Czechoslovakia,

German Democratic Republic and Federal Republic of Germany, Italy, Belgium and England, France, Switzerland, Austria, and Rumania (Pidoplichko, 1951; Krott, 1959) (V.H.).
Poland;

Geographic Variation

The wolverine
in

is

referred to as one of the

few mammalian species

and
in the

which, with a vast range, reveal insignificant geographic vari-

ation. This variation bears a clinal character,

Old World,

differences are expressed not so

much

in variation in meristic fea-

tures or qualitative structural differences of the skull as in color

variation.

the lightening of color

Geographic variation of this character is expressed in from west to east. This lightening occurs,

to another, but

however, not by means of gradual lightening from one population through changes in relative constituents of the sepaIn this way, wolverines in the west are darker

rate color types described above,

and of intermediate between them. "dark-brownish"

found undeveloped and extremely light

prevails, while others are
in

in

which the breast-band

is

almost

i.e.

"light-brownish" are few; those

which the saddle

is in

the

absent or extremely rare. In

form of a small spot are completely the extreme east, the picture is the

941
reverse, and in the

whole intervening extent

is

a region where, an

increase in

number of

light variants is observed. In general, geo-

graphic variation of wolverine, in particular the ratio of colored

types in different populations,
is

poorly studied.

Within the boundaries of our country, it is possible without doubt to distinguish 2 extreme forms. Probably, a third intermediate type may also be distinguished. 1. European wolverine, G. g. gulo Linnaeus, 1758. (syn. vul-

garis, borealis, arcticus, arctos).

Dimensions relatively small. Dark and very dark individuals predominate

rior
rare.

in

population.
Its

Breast-band not broad, dark or moderate color intensity.

ante-

ends do not unite on shoulders. Light individuals absent or

Condylobasal length of male skull 132.7-151.3 mm, of females, 128.7-148.2 mm; zygomatic arch of males, 92.7-110.0 mm, of females, 89.0-100.0 mm; interorbital width of males, 38.7-43.0

mm,

of females, 35.0-42.0

mm*;

length of upper toothrow of males,

51.0-56.3

mm,

of females, 49.0 to 52.0

mm

(Stroganov, 1962).
in

European part of country and in West Siberia. Borders relation to following form unknown.
Outside the
2.

USSR Scandinavian
g.

peninsula.
sibiricus Pallas, 1780 (syn.

East Siberian wolverine, G.

biedermanni, wachei).

Dimensions

as in preceding form.

Dark individuals and those of moderate color intensity predominate in population. Light forms occur more often than in
west. Breast-band quite broad and light.

Extreme

light individuals

absent or rare.

Eastern Siberia and Far East, except region occupied by fol-

lowing form. Borders

629

in relation to

Mongolian Repuband northeastern China (former Manchuria). Note: This form is conditionally given here for the first time. Possibly it is identical with the European which, however, is doubtOutside the
the northern part of
lic

USSR in

both other forms not clear.

ful.

The name given by

Pallas (1780)

may

possibly not belong to

the East Siberian wolverine. If the type locality of the

name

sibiricus

*In Russian original, erroneously given as "89.0 to 93.0"

Sci. Ed.

942.

is

considered 'upper

Ob"

i.e.

Altai, then

it is

not excluded that

it

it

belongs in the synonymy of the nominal race, since

that
it,

is

possible

and not the eastern form, lives in Altai. In order not to it probably is rational to transpose the type of form sibiricus locality the to the east (for example, Cis-Baikal). In any case, the names by Matchie as synonyms are related to the Pallas name. 3. Kamchatka wolverine, G. g. albus Kerr, 1792 (syn.
create

new names,

kamtschaticus Dyb., kamtschaticus Averin),

Dimension on average somewhat greater than in European form. Light and very light individuals predominate in population. Anterior ends of breast-band usually unite on shoulders, and whole band forms closed ring. Breech-band is light, often so broad that saddle is in form of small dark spot which is completely surrounded by light area of breast-band. Color of breast-band often light strawyellow and whitish or almost dirty white. Condylobasal length in males is 146.0-158.0 mm, of females, 142.0-149.0 mm; zygomatic width in males, 101.0-108.0 mm, of
females, 98.0-103.0

mm;

interorbital width in males,

41.9-44.2

mm,

of females, 98.0-103.0

mm;

length of upper toothrow in males,

54.5-59.7
1956).

mm,

of females, 54.2-56.8

mm (ZMMU

and Stroganov,

in Kamchatka and extreme northeastern Siberia (Anadyi Chukotsk and Koryakland). Absent outside the USSR. The details of the distribution of this form, the range of which is, probably, not restricted to the above-mentioned part of the northeastern extremities of Asia, and its limits to the west and south, are still unknown. The wolverine of the southern part of the Far East is darker, and seems not to be attributable to it. The Kamchatka form is a well-defined subspecies.

Found

Territory,

In

America
with

until recently, the existence of

cies

four

subspecies

has

been

one wolverine speaccepted (luscus,

katschemakensis, luteus, vancouverensis; Hall and Kelson, 1959).

However, a special study showed (Kurten and Rausch, 1959) that this Eurasian species is represented by one form only, G. g. luscus
Linnaeus, 1758.

943
primarily that the American form, craniologically well

It

is

differentiated
lar,

from the contemporary European one,

is

very simi-

apparently identical, with the form of the European Pleistocene.

that of the

Evidently, the rate of evolution of the Eurasian wolverine was

more intensive than

American (V.H.).

Biology
Population. The wolverine
is

a rare animal, never forming high

630

its relative numbers are supby information on prepared skins of hides: 70% of the catch Urals and 10% European North. Siberia and Far East, 20% However, the quantity of skins is a poor indicator; capturing this animal is difficult and sporadic. The typical density (per 100 hectares [= 1 km^]) of the wolverine in regions where this animal is common, fluctuates between 0.007 and 0.22. Densities of about 0.1-0.2 always are associated with high concentrations of ungulates. In British Columbia, 0.07 wolverine tracks were found per 10 km of route (Quick, 1953). One wolverine was caught on average in each 3.58 km^ (MacTaggert Cowan, 1957). In Sweden and Lapland (forest), the natural density in area with low yield of food was 0.01 [per km^] (two wolverines per 200 thousand

densities anywhere. Indications of

plied

hectares) (Krott,

1959).

On
the

the basis of approximate calculations of density and areas,

in the

number of wolverines
Habitat.

USSR may

be determined as 7,000-

7,500 individuals.

The wolverine
it is

fines of coniferous taiga

is widely distributed within the conand forest-tundra. It also enters the open

tundra, but here

rare. It

zone of the mixed forests

It is

much more rarely penetrates into the and to the West Siberian forest-steppe.
and
it

encountered both
its
it

in the plains

in

montane conditions.

Toward
although

landscape surroundings,
has been noted that

is

quite undemanding,

areas (Krott, 1959).

it is somewhat attracted to marshy Deep snow cover usually provides the wolver-

ine with an advantage in following

its

prey, and, therefore, the

main part of its range lies 70 cm and more; however it is not restricted therein. The winter period of deep snow (February-April) is particularly favorable to it (Teplov, 1955). The duration of the snowy period (220-240
days)

within the boundaries of deep snow; 50-

may

also be considered a favorable factor (Krott, 1959).

944
In
in

summer,

a time of

more food

security, the wolverine lives

very different habitats. Before the young

become independent,

the females are, probably, considerably less mobile than in winter

and do not go
In winter,

far

from the den.

in the

and

period of calving of ungulates, the wolsettlements. In Lapland, however, dens

verine confines itself to the region of their concentration. In Altai,

wolverine avoids
with
litters

human

located 2.5

km

from

a village

have been noted (Krott,

1959).

ent in

Food. The food composition of wolverine is essentially differsnowy and snowless periods of the year, a fact which was

established in several free-living tame animals (Krott, 1959). In

spring, they very intensively searched for and successfully obtained egg clutches of ground-nesting birds and destroyed them in great quantities. These were more often the clutches of ducks and other birds, and considerably more rarely, those of tetraonids (from 19481956, there were only 6 cases); this, apparently, was explained by

the difficulty the wolverines had in finding them.

They

also did

not destroy the crane nests found

in their area of activity.

wolverines

The larvae of wasps are of great importance in the nutrition of in summer. With the ripening of fruits, they give greater

attention to cloudberry, raspberry, blueberry, cowberry, cranberry

and crowberry. They rarely eat bog billberry. The wolverine prefers bird eggs, berries and wasp larvae to mouse-like rodents. The
latter

become,

in

"lemming"

years, of great significance in the

Scandinavian Peninsula. Cases of wolverine feeding on fish, amphibians, reptiles and crawfishes have not been noted; however, on
the Pechora,
cast
it
it

feeds on the carcasses of spawned-out salmonids

It is

up on the bank.

only rarely

that, in the

snowless period,

hunts hazelhen, capercaillie and white hares. The young animals

do not pursue forest game. They prefer carrion to fresh meat in summer. In summer, the wolverine does not attack ungulates and
small domesticated livestock, or does so very rarely (Krott, 1959).

Ungulates everywhere constitute the basis of winter nutrition

of wolverine, which
they locally
is

fairly well studied.

Most

frequently, these

are reindeer and elk. In the

montane

taiga of Siberia, evidently

become musk

deer, maral and

Manchurian wapiti, roe

deer and Siberian ibex. In the Altai (Dul'keit, 1953), in Lapland

(Nasimovich, 1948; Semenov-Tyan-Shanskii, 1948) and in other places, the wolverine feeds on ungulates mainly in the form of


945
carrion.

Within the Altai preserve, carrion constitutes 66.7% of

is

occurrences. In Pechora preserve (Teplov, 1955), with elk and

reindeer numerous, feeding on live animals
631

much more

frequent.

All ungulates in the food of wolverine constitute from 45 to

75%
of

of occurrences. Everywhere, the victims of wolverine are,

all,

first

those weakened from various causes, diseased and injured

its

animals as well as juveniles. Reindeer become

victims notably

more

often than elk.

Among

the latter, the

young more frequently

die (Teplov, 1955).

appearance

in

The share of carrion sharply increases with the regions inhabited by wolverine, of wolves and with

increase in their activity (Nasimovich, 1948). In the absence of the

latter together

with scarcity of ungulates, as well as in summer, the

role of ungulates considerably decreases

and the basic foods (20%

occurrence and higher) become white hares, tetraonid birds, forest

voles,

and

in

some

places nut pine "nutlets".

Among

those foods

of secondary importance are the squirrel, flying squirrel, fox and

others,

which are usually rarely

utilized in the diet.

It is

not a rare

weaker predators otters, pine martens and foxes. The latter are most often attacked in lairs while sleeping, sometimes near carrion. Not rarely, wolverine utilize the food remains of not only wolf, but also brown bear (Lapland) and also lynx (Dul'keit, 1953; Teplov, 1955). In Norwegian Lapland, in April 1951, remains of white hare, reindeer and willow ptarmigan occurred in a snow burrow where were found a litter of wolverines (Boehm, 1953).
event
the wolverine attacks as prey the

when

Wolverine, systematically though not often, rob the traps of

hunters eating the bait and the catch.

When

possible, the wolverine

it

plunders stocks in the hunter's hut. In rare cases,

eats fish

at

the expense of the river otter's catch, and getting the dead fish

thrown on the bank.

The wolverine's stomach can hold up

tuting about

to

2 kg of meat, consti-

17%

of

its

live weight.

It

was, however, shown that

feeding once a day, wolverine cannot eat more than 800-850 g, this is actually the maximum weight of its stomach content, i.e. it

does not consume the

is

maximum volume

of

its

stomach (Teplov,

1955). Therefore, current opinion about the voracity of this animal

eat "to satiety",

Even with abundant food, the wolverine does not and makes significant stores by pulling off pieces of meat and hiding them in the snow or at a tree in an area of up to 10 hectares around the animal carcass. Even in Pechora, where
not justified.

946
the wolverine has a good supply of food and is often distinguished by being well-nourished, it does not eat each day: from data on 12 daily tracks of wolverine, this was noticed in 7 cases (Teplov, 1955),

Home
eral years

range. Observations in nature during the course of sev-

on tame free-living wolverines in Sweden and Finnish Lapland (Krott, 1959) showed that each wolverine separating from
its litter,

takes over a vast, but quite definite

home

range or hunting

territory.

Within

its

boundaries

it

leaves

its

scent marks on logs, stones

and other noticeable places

(urine, secretions of the precaudal

and

abdominal glands). Between home ranges, there are no neutral zones. Therefore, despite very great daily movement, the wolverine is not
nomadic, but
within
its

is

a settled animal. In Swedish Lapland, one male lived

hunting territory for a period of not less than six years.

The area of such a hunting territory in the places with little is very great from 200-300 to 1000-1600 km^. In Swedish Lapland, the area of the hunting territory of an adult male was about 2000 km^ Exploring females occupied a hunting territory with an area 400-500 and 400-450 km^, and an exploring young male 700 km^. In areas with greater food, rich in ungulates and
food

tetraonid birds in Finnish Lapland, the hunting territories of wol-

verines were smaller

distance between brood dens

about 300 km^ females and maximum an average of about 20 km. In

for British

Columbia, the home range of wolverines equaled 124-132 km^

(Quick, 1953).

The dimensions of the hunting territory, therefore, are variable and depend on a series of features at the given locality. On the
other hand, the structure of the wolverine hunting territory
632 stable
is

very

a large hunting territory

for a male includes 2-3 smaller

female

territories.

Wolverines usually do not transgress on foreign hunting

territories.

On

penetrating a foreign hunting territory, they are

actively pursued and driven away, and fights occur

between males.

This aggression

is

characteristic of adult males and does not

extend to females (Krott, 1959). With predominance of carrion

feeding, the exclusivity of separate hunting territories
out. In several
is

smoothed
it

days large carrion sometimes attracts to

It

several

wolverines (for example, 3 wolverines in 25 km^).

that this occurs

is

possible

among animals from one

(1

litter

(cases were noted

when

the

litter

remains with the mother for the winter) or a group

male, 2-3 females).

forming a structural population unit

947

Using for the most part one and the same routes, often by

its

own
of

old tracks, the wolverine periodically visits the particular parts

In winter, these are mainly places of elk concentraetc.

its territory.

tion, the

winter pastures of reindeer,

Burrows and
nent shelter.
Its

shelters. In winter, the

lairs

wolverine has no perma-

usually occur under the protection of the

spruce crowns bent

places.

down

low, and more rarely on hidden elevated

its lair

The animal

prefers for

the places without an open

approach.
in

On

the Pechora, in the severe frosts, the wolverine hides

semiburrows in the snow. The dimensions of the lair are usually about 40 X 55 cm. For its litter of young, the wolverine makes a den differently: in a shallow ground burrow, under a twisted, downfallen tree or
directly in the

snow under dry

such a den,
is
it

Sometimes,

in

other cases, the lair

the fallen tree.

snow depth of about 1.5 m. makes a deep path in the snow, in protected from above only by the trunk of
logs in a
itself

The nest

may be
In

lined with dry fur and grass

localities or

or branches of spruce and

seashore, wolverine

fir.

montane

on the

makes

the brood den usually in a crevice

among

rocks or in the small caves in the very isolated places which are

young wolverines was found in Norwegian Lapland, in a snow den. The nest chamber was located 20 m from the entrance to the den. Daily activity and behavior. In the upper Pechora, the wolverine is mainly a nocturnal animal. Only in spring (March-April),
difficult to reach. In April 1951, a litter of three
i.e.

in the period of nursing the

young, diurnal activity

is

observed

more often
months,

up

to

75%

of observations

while

in

the winter

25%

(Teplov, 1955).

Lapland preserve, it is active in the period of polar night mainly during the day (Nasimovich, 1948); the observations carried out for several years in the Scandinavian Peninsula (Krott, 1959) led to the conclusion that a definite diel rhythm of
In winter, in
activity for the wolverine
is

absent.
itself from by ambush (sometimes from tree

Wolverine obtains
its

its

food by thoroughly concealing

pursuit,

victims,

by extended

branches), taking

away

the prey of other predators, eating the reIt

mains of
in

their meals,

and carrion.

waits for hare on the paths and

attacks tetraonid birds while they are resting, in mating places and

snow

holes. In searching for food, wolverine seeks out the tracks

it

of fox, lynx and others, but

avoids walking along wolf tracks. In

948
pestilence years, wolverine search out and eat carcasses of small

animals

hares, squirrels

and others. In

its

winter diel pattern

constitute up to

it

conducts
of

its

movements

in a straightline fashion,

mainly between

concentrations of ungulates. These

its

movements

54%

daily activity.

633

food and while fattening, its path is usually While walking, the animal examines large trees and the hollows in them, and looks under logs and stones. It willingly uses the old tracks of elk and ski-tracks of hunters to assist its movements on deep soft snow. In Sweden and Finnish Lapland, wolverines usually periodically examine their vast hunting territory covering a circle of about 1 km diameter. The outlines of the daily movement in these places usually have a serpentine appearance, with wide loops. Tracking a wolverine there is very difficult, particularly if it is followed, since it prefers to walk through "strong",
In searching for

twisting.

difficult of access, places (Krott, 1959).

and has great endurance. 8 to 45 km, averaging is not the maximum figure (Nasimovich, 1948; Teplov, 1955). In British Columbia, a wolverine traveled in one day about 32 km in one direction. In Swedish Lapland, an 8-month old male once went 34 km during the night. Following prey, a wolverine may move a distance of about 50 km. A case was also recorded when an adult male went 70 km without rest (Krott, 1959). The wolverine is well adapted to snow cover the weight load on 1 cm^of its foot (Teplov, 1955) ranges from 19.5 to 29.0 gm,
is

The wolverine

a very active animal

it

During a winter 21 km, but this

diel period

moves from

averaging 22 gm, and according to other data (Dul'keit, 1953),

27-35 gm.
It

climbs trees quite well, sometimes descending head downit

wards, but

never jumps from one tree to another (Krott, 1959).

Unsuccessful hunts constitute a

ine, especially

are

common phenomenon for wolverwhen pursuing ungulates. Seasonal migrations and transgressions. Regular migrations unknown in the wolverine. Rare cases of the wolverine's
in

appearance

the belt of

mixed

forests,

in the forest-steppe

of

West

Siberia (Vengerovsk and Kuibyshevsk regions of Novosibirsk

district)

and even

in

pine forests in the Kokchetav steppe,

its

may

in

some measure be

related to

eviction

beyond the

limits of its

range (Sludskii, 1953). Movements for a distance of 250

km

are

949

known

for separate individuals

(Naumov and Lavrov,

1948). In

years past, the appearance of wolverines in the Baltic region

Latvia and Estonia

were noted (Kh.I. Ling).

period from October to May. Apparently, they

Reproduction. In Pechora preserve, signs of estrus and mating

do not occur

in the

take place in the period June-September (Teplov, 1955). In

Mos-

cow Zoo,

signs of sexual excitement were observed in June-July

(Manteifel', 1947). In Pechora, parturition proceeds in the last days

of February and in the beginning of March; on Kola Peninsula

and the beginning of May. Therefore, the time of birth of the young for example, extends for two months, and pregnancy has a latent period. The investigations of wolverine reproduction carried out in Alaska (Wright and Rausch,
(Nasimovich, 1948)
in April

1955) on material from 22 males and 11 females during the period from 1950-1953, showed that in the October-December period, only unimplanted blastocysts were observed, and in the ovaries inactive corpora lutea, the fact of migration of blastocysts from one horn of the uterus to the other was established. In January and the beginning of February, only implanted embryos and active corpora lutea were detected. In April, females revealed signs of lactation and placental scars in the uteri were observed. It was found that in males, testes and their accessories [glands] begin to function at the beginning of February, but activity, with sperms in the accessories was only from the beginning of April. Therefore, in Alaska the breeding period extends from April to October;
copulation immediately after parturition, as in the polecat,
ologically impossible.
is

physi-

Observations from foreign zoos in various years showed that

copulation (lasting up to 30 minutes) has a local, variable form
the end of April, the end of

and the end of June (Krott, 1959). In Pechora, the time of birth of the young is the last days of February the beginning of March; in the Scandinavian Peninsula,

May

litter

found

in

February was demonstrated (Boehm, 1953). Ac-

litters from Sweden and Finland, within February-March, the age of the separate litters may differ by several weeks. The earliest case of parturition was on 11 February

cording to data from 13

(Krott, 1959).

A
i.e.

female having offspring remains barren the following year;

is

barrenness in adult females

not less than 50%. This also

litter

634 occurred in cases

when hunters

take the

from

her.

At Pechora,

950
the percentage of barren females is more than 60%. In Copenhagen Zoo, a female gave birth to offspring in two successive years (Wright and Rausch, 1955), which may be explained by conditions pertaining to captivity with a year round abundance of food. The litter usually comprises 2-3 young, rarely four. Out of 13 3 cubs each. cases, in 10 there were two young each, and in three Out of 38 cases, litters of three occurred 17 times, two 18 times, 2 times, and one 1 time (Krott, 1959). Average litter size four

in the

Scandinavian Peninsula
litters,

is 2.5.

In Pechora, according to data

most often 2 young occur (Teplov, 1955). In Alaska, judging from limited material (7 females), wolverine fertility was apparently higher, there were four corpora lutea in six cases and 5 in one. Parallel to them, there were 4 and 3 placental scars, 3 and 4 embryos, and [an average of] 4.4 and 2 blastocysts. Therefore the average number was revealed as 3.4; at the same time the difference in the number of corpora lutea shows an embryonic mortality equal to 17%; moreover, dead and resorbing embryos were observed. Besides the main den (see above), the female earlier prepares several reserve lairs, to which it transfers the litter in case of danger. The male never approaches the litter's place (Krott, 1959). In tamed animals, the first estrus was in three-year old females, but without subsequent pregnancy. Complete sexual matufrom 7
rity

appears simultaneously in both sexes at the age of four years

1959). In Alaska, sexual maturity of wolverine begins,

(Krott,

apparently, at the age of two years.

Among immature

males,

it

was

impossible to distinguish morphologically more than one age class.

from those sexually mature by the ermine, the growth of the latter character is due to effect of sex hormones; Wright, 1950). At the same time, the growth and development of the skull of male young

They

are easily differentiated

dimensions of the baculum

(in

wolverines take place very quickly

of
life

in the

course of the

first

year

(Wright and Rausch, 1955).

ratio in the population

Data on the sex

ity in
it

and the

rate

of mortal-

no data on its in February, had among In Alaska, one female, annual growth rate. 4 embryos, 2 males and 2 females (1:1). Among 22 captured males, 9 were adult and 13 were young; among 11 females 7 adults and
are almost absent, and, hence, there are

4 young, i.e. adult animals constituted 48% and young 52% (Wright and Rausch, 1955). Deriving from 3.4 corpora lutea per

951
adult female, the potential growth must be 75.5%, and embryonic

and postembryonic mortality

23.5%. Barrenness 83

in adult

females

(50-60%) lowers the calculated population growth two-fold, to 33%, and taking into account calculation of mortality to 25.2%. Growth, development, and molt. The weight of embryos sev-

eral

days before parturition equals 73 (oo)

gm

(o o) vvith

body length 122-125 mm. Eyes open, apparently, at the age of about 30 days. Lactation lasts about 3 months and, on Pechora, proceeds mainly during the period of deep snow cover (Teplov, 1955). In a series of cases, tracks of a male were noted around a
who, apparently, took part in the feeding of the young. On the Scandinavian Peninsula, no signs of male participation in the nourishment of young were observed (Krott, 1959). In
den with a
litter,

the northern Urals, already in early July, the litter begins to lead a wandering

way

of

life,

and by the beginning of winter, they

hardly differ from adults in size (Flerov, 1933).

In wolverines in captivity or

tame animals

in nature,

of the young to their attainment of sexual maturity,

to distinguish 5

it is

from birth proposed

phases (Krott, 1959).

The first phase is the first 4 weeks of life. Weight of males is 400-630 gm, of females 300-480 gm. In this time, they are still

blind, bear the first juvenile pelage

a dirty-grayish-yellow tone.

635

They drink their mother's milk exclusively. The second phase is the 4th-10th week after birth. During this period, the weight of young wolverines reaches 3.3 kg. Lactation continues. The eyes open in the 5th week, and by the 8th week, males are 10% heavier than females. The first juvenile fur is replaced by the second

a dark-brownish
is
is

color.

The

third phase includes age of

10-24 weeks. At the age of 11

weeks, average male weight

differs

3.8 kg, of female, 3.5 kg. In this

replaced by the summer fur, but somewhat from that of adults and resembles the summer fur of pine martens. At the age of 20-24 weeks, the animals eat 400-500 gm of meat per day. Males begin to exhibit more activity than females. The fourth phase includes the age of 6-12 months. At the beginning of August, the young wolverines don their winter fur. They begin to pursue live prey, but of dimensions no larger than
period, the second juvenile fur

themselves.

The fifth phase lasts from the age of 12 months to ment of sexual maturity. From the middle of April,

the attain-

the

young

952
wolverines already have acquired their second summer fur, not from that of adults, and in mid- August of the second year
life,

differing

of

the winter fur of the adult animal.

Summer marks
They make

the first

cases of preying on hazelhens, but usually the wolverine's attention

is

not attracted by tetraonid birds.

their first

attempts to attack small livestock, without killing the victims. With

the arrival of the second winter, a female tore apart a roe deer, and

young experimental male made an attempt to attack a small elk. one in the 3rd year of life killed six sheep in one night. Enemies, diseases, parasites, mortality, competitors, and population dynamics. With the exception of man and wolves, the wolverine does not encounter any dangerous enemies within its basic range. For man, the wolverine is a difficult, occasional catch. In the Scandinavian peninsula, wolves destroy wolverines (Krott, 1959). Bear, lynx, red fox, Arctic fox and large eagles are dangerous for young in the den. When the young follow the mother, lynx and bear as well as wolves and man, are a danger to them. On
a

In July,

being pursued by wolves, the wolverine jumps, if it is possible, into a tree. The damage caused by wolves must not be great, since
in the taiga

they [wolves] are rare, and the wolverine rarely trans-

gresses onto the tundra (Krott, 1959).

All carnivorous animals, especially those ranging in size from

red fox and lynx to bear

may

be, to a certain extent, considered as

competitors of wolverines. But factually, carrion and remains devourers such as these which utilize the remains from the table of
other carnivores are considered as food suppliers rather than
petitors. In all events, with the

com-

presence of wolves

in the region,

however, their feeding regime

is

obviously improved and second-

ary and occasional food use decreases.

The diseases and

ied.

parasites of wolverine

have been poorly studthe latter (47 to

In

the upper Pechora,

21%

of the wolverines caught were

infected with nematodes.

significant

number of

243 individuals) greatly weaken the wolverine. In some cases, pathological changes of the internal organs and exhaustion are connected with physical defects (the loss of feet, etc.). Longevity
is

unknown. Concerning fluctuations

in

numbers of wolverines, data are

limited. In Pechora preserve, the number of tracks encountered over an 11 -year period ranged from 0.02 to 0.04 along a 10 km route; i.e. two-fold. It was noted that the number of wolverines

953
increases parallel to the growth of the ungulate population in a

given region.
Field characteristics.

On

the

snow

surface, the wolverine leaves

well-defined prints of

all its feet,

with large well marked claws. In

it is

dimensions and

its

generally oval outline,

reminiscent of the

broad palm of an adult human.

The prints, which usually follow one another are almost always well distinguished because, even on the soft snow, the animal does not sink deeply. Walking in a straight line for a considerable distance is very typical for the wolverine. Excrement is rarely found along the track, usually not more than three times during a day's walk (P.Yu.).
636

Practical Significance

The

practical significance of wolverine is contradictory.

it is

hand,

a fur-bearing animal that provides a durable,

it

On one warm and

quite beautiful skin, though not expensive, and

taiga

sanitizes the

and tundra, destroying carrion and diseased and dying ungulates. On the other hand, with an increase in wolverine numbers, they begin to attack healthy animals, especially the young, and therefore in deer raising regions, they must be considered dangerous. In
forest

Sweden, the wolverine

since
it

is

considered a useful animal for the

in the

economy

reduces

number of ungulates
is,

in areas in

impossible for hunters to reach. The number of ungulates

cases, very great

many

and causes losses

in the forest (Krott, 1959).

Wolverine systematically steals bait and lures from the traps of commercial hunters and steals stores from the hunter's cabins;
therefore, they usually consider
it

an animal deserving of destrucis

tion the year round. In practice, this

difficult to realize. In the

State preserves, if
limited.

its

numbers increase

significantly,

it

must be

relative contribution in the

The world catch of wolverine comprises about 6,000 skins. Its USSR constitutes about 30%, and about 25% of the approximate number of animals in the USSR.
Wolverine is a very cautious animal, and different traps, in jawed traps, take them poorly. Setting the traps around carrion give somewhat better results. For hunting with dogs, ferocious and strong laika dogs are required, since the wolverine fiercely defends itself. Wolverine is very resistant to the action of poisons.
particular

954
All methods for
its

capture are complicated by the vast area of

its

home range and

searching out

its

very long daily movement.

is

In the Scandinavian Peninsula, the best effect

litters,

attained

by

which

is

carried out in the second half of or three hunters on skis

a typical area for fresh

March and the first half of April. Two with a 2-3-week stock of food search
still

tracks of a female leading to a den. In March,

blind, the female almost

when

the

young

are

always occurs near the brood. She is first killed, and then the live young are taken. Abroad, the zoos pay 250 dollars for young wolverines 12-14 weeks of age. In Sweden, the adult male hide is valued at 100 dollars, the female

70. Moreover, the hunters receive a bounty:

adult wolverine and

40

for young.

80 dollars for an Deer breeder organizations also pay

bounties for killed or captured wolverines (Krott, 1959) (P.Yu.).

Genus of Weasels and Polecats

Genus Mustela Linnaeus, 1758

1758. Mustela. Linnaeus. Syst. Nat., ed. X,
1,

p.

45. Mustela

erminea Linnaeus.
1775. Putorius. Frisch. Natur-System 1817. Putorius. Cuvier. Regne anim.,
d. vierfUss. Thiere, p.
1, p.

11.

Linnaeus.

Entw. Gesch. Nat. Syst. Europ. Thierw. 1, Mustela erminea Linnaeus. 1829. Ictis. Ibidem. 1, p. 35, 40, 41. Mustela vulgaris Erxleben = Mustela nivalis Linnaeus.
1829. Arctogale.
p.

30.

. .
1,

147. Mustela putorius

1840. Foetorius. Keyserling

et Blasius.

Wirbelthiere Europas.

p.

68. Mustela putorius Linnaeus.

1841. Gale. Wagner. Schreb. Saugeth. Suppl.

vulgaris Erxleben

2, p.

234. Mustela

= Mustela
2, p.

nivalis Linnaeus.

1841. Lutreola. Wagner. Ibidem.

637

239. Viverra lutreola Linnaeus.

1871. Mustelina Bogdanov. Birds and animals of the chernozem

belt of

Povolozh'e

(Tr.

Obshch. estestvoisp.

pri.

Imp. Kazansk.

Universit.,

section

1,

p. 167. Mustela erminea Linnaeus*.

The mink was incorrectly referred to as the type of this genus by Ellerman and Morrison-Scott (1951), as well as the type of the following Bogdanov genus (which
is

correct).

955
1871. Hydromustela. Bogdanov. Ibidem,
167. Mustela lutreola

p.

Linnaeus.
1899. Eumustela. Acloque. Faune de France,

Mamm.,

p. 62.

Mustela

vulgaris Erxleben (= Mustela nivalis Linnaeus) et Mustela

erminea Linnaeus.
\9\\. Kolonokus. Satunin. Izv. Kavkazsk. muzeya, 5, p. 264.

Mustela sibirica Pallas (V.H.).

General dimensions small or very small. The smallest species
of the order belong to this genus.

The

skull is usually relatively small, flattened in the majority,

strongly elongated and narrow, without crests and protuberances.

The braincase

is

voluminous and elongated,

is

slightly

widened

in

the occipital region, the facial part

short,

weak and

blunt,

zygomatic arches are

thin,

weakly separated (often not as wide or

just wider than the cranial part of the skull). Supraorbital proc-

esses are small, the interorbital and postorbital constrictions are

weak
fined.

or very

weak

or else the postorbital process

is

is

barely de-

The

dorsal profile of the skull

usually straight and the

convexity in the orbital region not developed or weakly developed;

the profile of the facial portion
is

only slightly and gradually deis

pressed.

The diameter of
it

the infraorbital foramena

greater than

that of the alveolus of the

upper canines;

in

some

species, they are

equal to
In a
is

or even less.

few forms, the whole skull is wider and shorter, the brainand the facial part is strengthened, the zygomatic arches are relatively robust and more widely separated, the postorbital constriction is relatively well or even strongly
case
relatively shortened

defined, the convexity of the upper skull surface in the interorbital

area

is

stronger, crests and protuberances, etc. are quite sharply

developed.

Some

species have skulls of an intermediate character.

in all species are

The auditory bullae

situated

notably swollen through-

out the lower surface, usually of elongated bean-like form, and

more or

less parallel to

each other;
anterior.

in

some

species, they

are relatively shortened and the posterior portions are relatively

more widely divergent than the

The hook-like processes

latis

of the pterygoid bones are not united with the auditory bullae. The

mammary

(mastoid) processes are weakly developed, and the

eral occipital (paroccipital) are not developed.

The bony

palate

of moderate width. In general, with a comparatively large

number

956
of species, several of them have evident differences in skull structure,

though

in close species, they are not great (see later).

Dental formula

I-C-P M = 34.

13

There
last

is

no additional cusp

on the inner side of the main crest of the

tooth behind canine

lower premolar (third

The lower carno additional cusp on the inner side of the median crest. The longitudinal axes of the crowns of the upper carnassial teeth lie at a significant angle to each other and with the longitudinal axis of the skull. The second upper premolar (first in the row) is very small, and correspond approximately in dimensions to the first [premolar] in martens (genus Martes), but
^just

anterior to carnassial).

nassial tooth (first molar) has

it

is

not lost or this occurs only rarely.

The trunk

is

thin

and strongly elongated,

it

in

some

species to an

extreme degree, and

appears as though "snake-like". The legs

tail is

are short, in several, very short, the

of various lengths and

may
it is

attain half or nearly half the

body

length. In several species

less

short,

and constitutes about 0.25 of the body length or

is

and

perhaps equal the length of the hind foot. The head

638

relatively

small, narrow, elongated and flattened, usually no broader in diameter than the neck, in

some almost "snake-like"

are digitigrade;

in

appearance. The

ears are small, very widely separated, rounded; the eyes are quite
large.

The extremities

palms and soles are moderin all species,

ately broad.

Anal scent glands are developed

and

in

some

very strongly; individuals are capable of spraying the secretion of

these glands. All have a special "polecat" odor.

Abdominal glands

characteristic of wolverines and martens are absent.

The

fur

is

dense, but usually not long, and in the majority,

closely compacted; in some, quite fluffy, and

more

or less equal in
is

length over the whole body. In the majority, the fur

silky

soft

and
cov-

and

in

some, highly valued. In the majority, the

is

tail is

ered by short hairs and

relatively thin, but in

some

quite fluffy.

The color is quite variable monocolor ocherous or brown

dark venter.

two-toned with
in the

a light venter, of

tones, or blackish-yellow with a

distinct throat patch

does not occur. In some spe-

form of a "mask" of dark or, on the contrary, light color. In one species with monotone general color, there is a narrow white band along the whole back and on the belly (M. strigidorsa).
cies, there is a pattern

on the head

957
Seasonal dimorphism
ern forms
is

in density

very pronounced, while in the south,

and length of the fur of northit is weaker or

even entirely unexpressed or nearly so. In some northern species, sharp seasonal dimorphism in color is very pronounced (whitening in winter), the sharpness of its manifestation in general and even
the reality of change in color itself varies geographically within the

one species. Sexual differences in color and character of the fur are absent; in all species, females are smaller than males, in some quite significantly. There are 4 pairs of inguinal and ablimits of

dominal mammae, or

in several, 3.

All representatives of the genus are very energetic, quick, active

and clever animals. The majority are able to climb trees, however, there are arboreal forms in the genus, and all of them are true terrestrial carnivores, usually moving in leaps, bending the back. All climb very well in windfalls, among rocks and in rodent burfrom the Arctic and rows. They inhabit very diverse biotopes high mountains higher than 3000 m above sea level to steppes and

deserts and tropical forests. All of

them can swim

some species
reveal

are typically amphibious forms associated with fresh water (minks).

Some

species do not avoid proximity to

humans and even

some anthropophilous tendencies. They are settled forms; monogamous; to 10 young. Lairs are made in extremely
of rodents. They do not

there are 1-2 and 3-4,

varied places

among

windfalls, in rocks, etc., but chiefly in underground burrows, mainly

make

nests in tree hollows, especially high

above the ground. They are usually solitary animals, but several sometimes hunt in small groups (litters?). They are sharply distinctive carnivores, utilizing no plant food at all or almost none. The main prey are rodents of various sizes from mice and voles to large ground squirrels. The majority are miophagous*. Birds, reptiles
and amphibians (frogs) are foods of secondary importance. Sometimes there
is

a quite close connection with particular species of

rodents (the steppe polecat

weasel

small voles

ground

squirrels,

ermine

water

vole,

etc.).

Several forms are specialized for feed-

ing on fish as well as amphibians, crawfishes etc. Hibernation or

winter sleep does not occur.

The species of

the genus are quite variable in size.

The small-

est (weasels) have, in several

(averaging about 160

*Lit. "mouse-eaters".

mm)

forms a body length from 130 and a weight of less than 100

mm
gm

This seems to be a coined word

Sci. Ed.

958
(40-75

gm

the smallest species of the carnivore order),

and larg-

est species (M.

eversmanni)

[body

length] to 565

mm.

640

the genus is very vast and occupies the greater and the northern part of North Africa, all of North America and a considerable part of South [America]. The northern limit of the range in America includes the whole mainland and the entire Arctic archipelago and the northern and northeastern part of
part of Eurasia

The range of

Greenland south

to

approximately 70 N.

lat.

or a

little

southward.

The southern

limit passes along the northern

and northwestern part

of South America.

To

the south

it

covers Venezuela and south-

western Columbia, to the west includes Peru and to the west and
south the Bolivian Andes to the latitude of Lake Titicaca or a
little

southward, and passes to the Pacific Ocean. On the whole, on this continent, the representative of the genus (M. frenata) is mainly associated with montane regions where it attains a height of 3,000 m above sea level. Species of the genus are encountered on all of the islands of the Atlantic coast of North America and on all the
islands of the Pacific coast, including Kodiak, but are absent in the

West
St.

Indies.

On

the islands of the Bering Sea (Aleutians, Pribilofs,

Matthew, Diomedes and others), it is absent except on Unimak'. In the Old World, the range occupies all of Europe except Iceland, the Arctic Islands and the islands of the Mediterranean Sea. In Africa, the range includes its extreme northwestern corner^ Morocco, the parts of Algeria and Tunisia connected with the Atlas mountain system and Egypt, apparently, only Lower.

In Asia, the northern limit of the range occupies the entire

mainland and the

Iraq. In Iran, the
try,

New

Siberian Islands.

The southern

limit in Asia

includes, to the south, Palestine, Syria and the northern half of

range occupies the northwestern part of the coun-

whence,

it

passes as a broad projection along the Zagros

moun-

tain

system towards the southeast; however it does not extend far, not reaching the shores of the Persian Gulf (extreme southwest of the country is not included in the range). Farther on, the range
occupies northern and northeastern Iran (representatives of the genus
are absent over the entire central desert part),

apparently, to the south covers the

whence the border, Hindu Kush system, through

'References of several authors (Carter, Hill and Tate, 1946) on the occurrence of weasel and mink on the Aleutian Islands, are apparently, mistaken and are not confirmed by other authors (Miller and Kellogg, 1955; Hall and Kelson, 1959). On

Unimak,

the existence of weasel

and ermine was recorded (Murie, 1959).

960
Khazaradzhat (Hazara) and, apparently, the Kabul region and passes
to India

to Chitral.

Farther to the east, the range occupies Kashmir and the entire

Himalayas from Kashmir through Nepal, Sikkim, Bhutan, and to Assam inclusively. In southeast Asia, the range includes Burma, the Indochinese Peninsula, Tenasserim, Malacca and the islands of Sumatra, Java and Kalimantan (Borneo). Eastward the mainland range reaches the Pacific Ocean and includes the islands Karangin, Kuril (in part), Shantar, Sakhalin, the large Japanese [islands] (Hokkaido, Honshu, Shikoku, Kyushu) the Ryukyus, Taiwan and
genus are absent. The range of the genus has not suffered noticeable changes on the negative side under the influence of human activity in the time under review. The ranges of some individual species have someIslands, representatives of the

On Commander
Hainan.

the Arctic Islands, except

New

Siberian, as in the

times changed in recent times, either negatively or positively.

The scope accepted here of

the genus (13 species) itself rep-

The extreme members are a more or less natural series, which may comprise the species of the genus from the weasel, M. nivalis (races of the group rixosa) to the white polecat (M. eversmanni), differing from each other in many
resents a quite heterogeneous group.
respects,

among which

are the craniological, significantly.

On

the

other hand, in several characteristics, separate species within this

series are obviously similar to each other. This

gave an opportu-

nity to various authors to divide the

rate genera or subgenera. Thus, separate

genus accepted here into sepagroups were distinguished

such as polecats (Putorius), true weasels and ermines (Arctogale), minks (Lutreola), Siberian weasels (Kolonokus) and Javan kolonok (Plesiogale) in part, combining in different ways the species in
these groups.

At the same time,

whole,
641
it is

if

we

consider the group of species as a

it

sufficiently clear that strict delimitation of

into sev-

eral is impossible,

and the most extreme forms, as shown above, are connected with each other by a series of intermediates. This
applies
is

all

the

more

to other

members of

the genus. Therefore,

it

which two subgenera only may be distinguished the specialized polecats: subgenus Putorius and subgenus Mustela, to which belong all remaining species. Even separation into a subgenus of Siberian weasels and minks, which up to now has been accepted by some authors
natural to accept the large genus Mustela in

961

(Stroganov, 1962) cannot be considered well founded

considers
all

if

one
are

species of the genus.

The polecats even now

that has

been poorly received. The mink serves as a connecting link between them and the remaining species^.

sometimes separated into a separate genus {Putorius),

The
nus
is

identification of the actual systematic position of the ge-

among

these various forms (genera) present in the subfamily,

It

very difficult.

is

apparent, however, that this

is

a line of

development
wolverine.

in the

subfamily different from the true martens and

the other genera of the sub-

It is

more connected with

family (see above) than with Martes and Gulo ("Martinae" and

"Guloninae" of some authors). Despite the presence in it of some specialized forms (subgenus Putorius), in its group of genera the
genus Mustela must, apparently (in the pure systematic sense) be placed "in the foundation" of all groups of Mustelinae (except the

two above-mentioned genera). This

cal
(their specialization

is

indicated by the craniologi-

structure of the species belonging to the subgenus Mustela

obviously proceeded in another direction,

in

form of the body). The genus Mustela, apparently, is more ancient to it belong several forms from the upper Miocene of North America and Europe and from the lower Pliocene of Asia. The closeness of the connection between these fossil forms is not clear. The following 13 species may be included in the genus (listed nivalis in order of increasing specialization): weasel, M. separate American authors consider a which (including M. rixosa species); ermine, or suslennik* M. erminea; solongoi, M. altaica Indian ermine; M. kathiah (sometimes this species is united with M. altaica); kolonok, Siberian weasel, M. sibirica (including the form itatsi, sometimes considered a separate species; Stroganov, 1962); long-tailed weasel, M. frenata; white-striped weasel, M. strigidorsa; naked-footed kolonok, M. nudipes; European mink, M. lutreola; American mink, M. vison; black or forest polecat, M. putorius; American polecat [black-footed ferret], M. nigripes; white
particular the elaboration of a special

sian

^An indication is that hybrids may exist between the black polecat and the Rusmink in nature. For details on relationships in the genus, and connections between
*Local name referring to

the polecat group

on one hand and a remaining species see section on polecats and mink. suslik, or ground squirrel, which in many places is an

important prey species

Sci. Ed.


962
belong to subgenus Mustela. Of the 13 species, two are Eurasian-North American (M. erminea, M. nivalis, the latter also in North Africa), 1 is North and South American (M. frenata), 2 are North American {M. vison, M.
or steppe polecat,
last three species

M. eversmanni. The

the subgenus Putorius, and the remainder

to

nigripes), 2 are
in the

European (M. putorius, M.

lutreola, the latter also

Caucasus), 3 are South Asian (M. kathiah, M. strigidorsa,

M. nudipes), and 3 are Central Asian and Siberian {M. eversmanni, M. sibirica penetrating also into eastern Europe; M. altaica). They are fur-bearing species, and are destroyers of rodents and regulators of their number. The indigenous fauna of the USSR comprises 7 species: 1) weasel, M. (/.) nivalis Linnaeus, 1758; 2) ermine, M. (M.) erminea Linnaeus, 1758; 3) solongoi M. (M.) altaica Pallas, 1811; 4) Siberian weasel [kolonok], M. (M.) sibirica Pallas, 1773; 5) Russian mink, M. (M.) lutreola Linnaeus, 1761; 6) black or forest polecat, M. (P.) putorius Linnaeus, 1758; 7) white or steppe polecat, M. (P.) eversmanni Lesson, 1827. One species the American mink, M. (M.) vison Schreber, 1777 has been acclimatized. Species of the genus occupy the entire territory of the country and are encountered in all biotopes. They play an important role in fur trade, and in part as regulators of populations of rodents pests of agriculture and transmitters and reservoirs of dangerous diseases of humans and domestic animals (V.H.).

642

Key for
1

Identification of the Species in the

Genus Mustela

(4).

Color of entire body monotone white (winter) or lower

side of

body pure white (sometimes with

lip

light grayish-

yellow highlights), upper side various brownish or sandy

tones (summer). Color of upper
tiated

and chin not differen-

from

that

of surrounding parts of head. Color

borders on sides well marked. Skull moderately elongated

distance between mastoid

(mammary) processes approxiskull.

mately equal to half of condylobasal length of the

2(3).
Distal half or third of
tail

pure black. Tail length with

terminal hairs equal to half of body length or

somewhat

more. Width of skull below canines notably less than

width of interorbital area. Condylobasal length of male

963
skull
is

up

to

53

mm,

of female

up

to

46

mm
may be
tail

ermine,
3(2).
Tail

M.

(M.) erminea (page 995).

indi-

monotone along
its

entire length (there

vidual dark hairs at

extreme end). Length of

with

terminal hairs significantly less than half, usually less

than one-third, of body length. Width of skull below the

canines approximately equal to width of interorbital area.

Condylobasal length of male skull up

4(1).

to

47.2

mm;

of female

weasel, M. (M.) nivalis (page 965). up to 36.7 mm Color otherwise. If dorsum tawny and brown tones, and venter light and border between dorsum and venter colors

5(8).

on sides well marked, then distal half of tail not black and not sharply differentiated from color of basal portion and length of tail with terminal hairs about half to more one-third of body length^. Skull moderately or greatly elongated or relatively short and broad. Lower surface of body, except throat and chin and sometimes middle of abdomen, black or blackish, darker than dorsal side; legs always black. Entire tail or its distal half black. Skull short and broad distance between the mastoid processes considerably greater than half of

condylobasal length of skull.

6(7).

Dorsal body color dark with black color predominant, through which very light yellowish underfur shines
through, especially on sides; venter black or blackish. All
or almost
all

of

tail

(except very base) black. Black color

of legs not sharply contrasting with color of trunk.

Postorbital part of braincase has no well-defined isthmus

and
7(6).

its

edges form parallel or almost parallel lines

black polecat,

M.

(P.)

putorius (page 1107).

Color of upper body light with yellow color predominant, because light underfur is weakly covered by dark-brown
guard hairs; venter
light,

yellowish. Only distal half of

tail

black; basal part has light color of back. Black leg color

sharply contrasting with color of trunk. Postorbital part

of braincase with well-defined isthmus, and

its

edges

form angled
....

lines, corners directed

towards each other

steppe (white) polecat,

M.

(P.)

eversmanni (page

1133).
'An exception
text.
is

present in

some

races of solongoi [M. altaica]; see 14 (13) and

964
8(5).

Color of abdomen and lower side of body not black or tail without black color, its color corresponds to upper body. Skull moderately elongated
blackish, legs not black, or extended (mastoid width of skull constitutes about half,

or less than half, condylobasal length of skull).

9(12).

Color of entire body monotone

dark tawny-brown with

dark undercoat; venter not lighter than dorsum. Small

white marks only on lips and chin, sometimes on chest.
643

Skull moderately elongated

distance between mastoid proc-

esses constitutes about half of condylobasal length of skull.

10(11). Both lower and upper

lips white. Least width of skull behind supraorbital processes greater than interorbital width or equal to it. Upper molars small, and longitudinal

diameter of

its

inner blade

is

a bit greater than outer.

Posterior end of second premolar in contact with extreme

antero-outer angle of carnassial tooth

...

Russian (European) mink,

lip

M.

(M.) lutreola (page 1078).

11 (10).

Only lower

white. Least width of skull behind

supraorbital processes less than interorbital width.

Upper

molar relatively large and longitudinal diameter of its inner blade markedly greater than diameter of outer. Posterior end of second premolar contacts carnassial tooth some-

12

(9).

what more mediad to antero-outer corner of carnassial American mink, M. (M.) vison (page 1391). Entire body color if not monotone is not dark tawnybrown, but bright yellowish red or dull yellowish-strawsandy, or below is somewhat lighter than above, or vice versa. Skull elongated distance between mastoid processes
tooth

is

usually less than half of condylobasal length of skull.

13 (14). Color of muzzle and area near eyes coffee-brown, forming well delineated dark
striction of skull

"mask" on head. Postorbital con-

elongated, with almost parallel outer

outlines and does not form sharp isthmus directly behind

supraorbital processes. Color bright yellowish-red or

brownish-red, lower side not lighter than upper or only

slightly lighter. Transition

between ventral and dorsal color

always completely gradual. Dimensions relatively large

(condylobasal length of skull of adult males more than 55

mm,

of females

more than 48 mm)

M.
(M.) sibirica (Page 1051).

Siberian weasel,

965
14(13). Dark "mask" on muzzle absent. Postorbital constriction
of skull short,
its

lateral outlines not parallel

and forms
little

a sharp isthmus directly behind supraorbital processes.

Color
to

is

bright or dull, yellowish-red, lower side a

considerably lighter than upper, perhaps almost dirty-

white with ocherous highlights, boundary between colors

of dorsum and venter sharp or transition between them
gradual. Dimensions smaller (condylobasal length of skull

of males less than 55

solongoi

mm, of M. (M.)

females

less than

48

mm)

altaica (page 1030) (V.H.).

Subgenus of Weasels
Subgenus Mustela Linnaeus, 1758

WEASEL'
Mustela {Mustela) nivalis Linnaeus, 1766
1766. Mustela nivalis Linnaeus. Syst. Nat., ed. XII,
1,

p.

69.

Westerbotten province, Sweden^.

644

1777. Mustela vulgaris Erxleben. Syst. Regni Anim.,

1,

p.

471.

Umerennaya Evropa [Temperate Europe]. After Ognev (1935) Vic. Leipzig.

1800. Mustela boccamela. Bechstein. Pennant. Ubers vierf. Thiere,
2, p.

395. Sardinia.
3, p. 94.

1811. Mustela gale. Pallas. Zoogr. Rosso-Asiat.

ing of vulgaris.

Renam-

1877. Mustela stoliczkana. Blanford.

p.

J.

Asiat. Soc. Bengal, 46, 2,

260. Yarkand.
p.

1896. Mustela rixosa Bangs. Proc. Biol. Soc. Washington, 10,

Saskachewan, Canada. 1899. Mustela nikolskii. Semenov. Zap. Akad. nauk,

21.
p.

ser. 8, vol. 6,

14. Vic. Simferopol, Crimea^.

this species is

^^2

^^^^ correct spelled name for

the verb lazat' [=to climb].

undoubtedly, lazka, and

is

derived from
agility,

The

ability
is

of the animal to climb with extraordinary

especially

among

fallen limbs etc.,

^Westerbotten province lies encountered references to the type locality, said to be in northern Sweden, are incorrect. This has substantive nomenclature significance. ^This form is incorrectly cited in Ellerman and Morrison-Scott (1951).

and immediately calls attention to itself. between 60 and 61 N. lat. and therefore, often
striking,

966
1900. Putorius nivalis pallidus. Barrett-Hamilton. Ann.
Hist. 5, p. 48.

Mag. Nat.
p. 48.

Kokand, Fergana.

1900. Putorius nivalis caucasicus. Barrett-Hamilton. Ibidem,

Caucasus, Gochal (?) mountains, 12000'.

1901. Mustela (Ictis) dombrowskii. Matschie. Sitzungsber. Ges.
naturf.-Freunde Berlin,
p.

231. Siul'nita. Romania.

J.

1903. Putorius (Arctogale) pygmaeus.

Sea.

Allen. Bull.

Amer. Mus.

Nat. Hist., 19, p. 176. Gizhiga, northern shore of Okhotsk

1907. Putorius nivalis dinniki. Satunin. Izv. Kavkazsk. muzeya,

p.

3,

105. Stavropol (Caucaskii).

191

1. Ictis

1921.

1922.

1926. 1927. 1936. 1936.

boccamela nikolskii. Satunin. Izv. Kavkazskovo muzeya, 257. Crimea^ Mustela rixosa namiyei. Kuroda. Journ. Mamm., 2, p. 209, Aomori, northern part of Honshu (Hondo) island, Japan. Ictis nivalis var. kamtschatica. Dybowski. Arch. Tov. Nauk. Lwow, 1, p. 349. Nomen nudum. Mustela punctata. Domaniewski. Ann. Mus. Zool. Polonici. N.H. 5, p. 55. Darasun; southern Trans-Baikaliya. Mustela nivalis mosanensis Mori. J. Chosen. N.H. Soc, 5, p. 28. lengan near Mosan, Korean Peninsula. Mustela pygmaea yesoidsuna. Kishida. Dobuts Zasshi, 48, 4, p. 177. Hokkaido, Japan. Mustela pygmaea karaftensis. Kishida. Ibidem, p. 177.
5, p.

Sakhalin.

1937. Mustela trettaui. Kleinschmidt. Falco, 33,

Isp. Prirody, 57,

p. 11.

"Germany".

1952. Mustela nivalis kerulenica. Bannikov. Byull. Mosk. Obsch.

No. 2, p. 40. Vic. Undurkhan on Kerulen Mongolian Republic. 1953. Mustela nivalis heptneri. MoToso\a-Turo\a. Zool. Zhurn., tributary of Kushka [river], 37, No. 6, p. 1267. Egri-Gek Turkmeniya Badkhyz; southern (V.H.).
[.],

Diagnosis
Coloration monotone white, or dorsum dark tawny tones, and
venter white, with color boundary between dorsum and venter

sharply demarcated. Distal half of

Satunin, apparently, not

tail

not black. Tail length

the

knowing of

the

work of Semenov, describes

Crimean

weasel as a

new

form.

967
significantly less than half of

body

length, usually not

more than
mas-

one
toid

third. Skull is

moderately elongated

distance between

(mammary) processes approximately equal

to half of condylobasal

length of skull.

to interorbital width.

Width of skull below canines approximately equal Dimensions small to very small (V.H.).
Description^

645

The general appearance of

the weasel

is

very characteristic.

It

has

body with a small, somewhat elongated head with a small, blunt muzzle. The head is
a thin, greatly elongated, extremely flexible

narrow, not thicker than the neck; the eyes are quite large, dark

and
is

slightly bulging; the ears are situated

wide

apart, short

and
tail

rounded. The legs are short, the feet of moderate breadth. The
relatively short.
It is

considerably less than half the body length,

its

as a rule not greater than one-third of

length,

sometimes not as

long or slightly longer than the hind foot, and in individual cases,

even shorter than the hind hairs, and appears thin, not

foot,

covered with short, close-lying

fluffy.

The lower surface of

the front

and hind are

feet

covered with hairs, the claws are not long, but

very sharp, dark.

The small head, long neck and thin, elongated body (where the head can go, so can the body) confers on the weasel, especially with its extreme mobility, a unique "serpentine" appearance. When moving, the weasel proceeds by jumping, strongly bending the
back; "creeping" very quickly and nimbly

among

rocks, brush-

wood, rodent burrows etc. Winter fur is dense, but short, and quite closely-fitting; in northern forms, it is soft and silky and in southerly forms, coarser. In summer it is very short, sparser and rougher. In northern form of the species, variations in characteristics of winter and summer
fur are considerable, but in southern forms,

much

less.

coloration

body including the and inner sides of the legs are white. Sometimes the palms and the lower parts of the anterior extremities and the toes of the posterior are partially or completely white. The upper is dark and in different races, varies greatly from dark-tawny or dark-chocolate to light pale tawny or sandy. The color of the dorsum extends to the
is

bi-colored

the lower

Summer, lower jaw

'In the case of characteristics of this species,

some unpublished

materials,

made

available by L.G. Morozova-Turova, were used.

968

645

Fig. 234. Weasel,

Mustela (Mustela) nivalis L. Sketch by A.N. Komarov.

outer side of the legs; the

tail

has the same color as the back. The

boundary between colors of the dorsum and venter on the sides is straight but sometimes forms an irregular line. There are sometimes dark-tawny spots on the white lower surface of the body, more often behind the corner of the mouth. The color in winter is pure white and only in the form of rare exceptions do individual dark (black) hairs occur on the very end of the tail, but a black tail tip is not formed. Only the most southerly forms
in

do not whiten

at all in winter, or

color

may

take place, or the side

only some lightening becomes white, but the white

color does not cover the middle of the back.

When

molting, the

animals sometimes become scattered over a white field.

however,
different
646

piebald

irregular

dark parts are

Individual differences in fur color are in general insignificant,

in various races, the

amplitude of these differences

is

and

in several

(Western Europe), these differences

may

be quite great. In particular, there is variation in the character of the boundary of coloration between the dorsum and venter and the frequency of appearance of dark marks on the light field, especially the

lower jaw. Age and sex differences

is,

in color are absent,

but geographic

as

shown, significant.

969

The

skull is moderately elongated, with a

voluminous, quite

broad and swollen brain case. Its anterior region (in the area of the frontal bones) is wide and voluminous, the postorbital constriction
is

well-defined, but short and not deep. Facial parts of the skull are
short.

above the canines is approximately equal to interorbital width. The zygomatic arches are moderately divergent (zygomatic width approximately corresponds to greatest width of the skull posteriorly), thin and weak. The infraorbital foramen is small; its transverse diameter is approxi-

weak and very

Width of

the skull

mately equal to the longitudinal diameter of the canine alveolus.

The upper
evation on
it.

profile of the cranial region of the skull

is

slightly

flattened and the interorbital area does not

form

a significant el-

Sculpturing of the skull

is

weak

the arrow-shaped

[sagittal] crest is not

defined or weakly defined in the posterior

part, the occipital is relatively

considerable.

and their

more strongly developed, but is not The auditory capsules are bean-like in form, convex inner edges are parallel. The distance between them is

approximately equal to the width of the hard palate at the level of the anterior border of the sphenopalatine notch. Teeth are relatively small, but the carnassial tooth
is

developed and the canines

are long and strong, although thin.

On the whole, the weasel skull, especially that of the smallest forms {rixosa group) has infantile features as compared with other closely related species of the genus (ermine, Siberian weasel). These
are expressed in the relatively large size of the entire cranial portion,
in

the size of

its

anterior part (on the extension of the

zygomatic arches

the

frontal part), in the unusual shortening of

the facial part, the

weak development of

the crests and in general

sculpturing. Apparently, here, as well as other likely reasons, the

phenomenon of allometry may

ters.

also be expressed.
is

Individual variation in the skull

considerable in

all

characis

The sexual

is

illustrated by the fact that the skull of males

in the

larger,
is

and wider

mastoid

part, the postorbital constriction

absolutely and relatively narrower, and crests, tubercules etc. of

the skull are

more strongly developed.

is

Age

variation of the skull

is
still

insignificant. In

young animals,

the brain case

of relatively greater size than in the adults

and is more swollen. It is characteristic that the sequence of age changes in the skull is small and take place so quickly that the skull of a young weasel in the first winter is only a little different

970

from

that of the adult.

The complete fusion of

the skull sutures

occurs relatively very early

the genus.

earlier than in the large

members of

Geographic variation of the

skull, in

accordance with the genIt is

erally large variation of the species, is very great.

expressed

both in general dimensions and also in general structure and proportions of

its parts.

In the small forms, infantile features are

more

strongly developed than in large forms. In the latter, in connection

with the strengthening of protuberances, crests and other features,

similarity to the ermine skull in particular increases.

The degree of

sexual dimorphism in dimensions changes geographically

smallest forms,
it

in the
in-

is

almost unexpressed, but grows with the

is

crease in general dimensions and

races.

well developed in the largest

is

The OS penis is sharply bent upwards at its distal end, and here formed a hook, characteristic of the majority of species of this The dimensions of weasel

genus.
individual variation, rarely found
in the race vulgaris,

the

some races to great among mammals in general. Thus, weight of the largest individuals may
are subjected in

exceed the weight of the smallest by almost four times (35 and 130 gm). In some large races males may be almost 1.5 times larger
than females (body length). Geographic variation in dimensions
648
is

very great, greater than that in other species of the family. The

average weight of animals of large races

may be 2-3

times more

than that of small races. Weight of individual animals of the large

may exceed the average weight of small races by 7-8 times. Not only absolute dimensions and weight vary geographically, but also the amplitude of their individual variation. Geographic variforms
ation in relative
tail

length

is

characteristic

in various races

it

constitutes

from 13

to

30%

of the body length. In the extreme case,

length
it.

in individual

animals the

tail

may be

less than that of the

hind foot length or equal to

130-260 mm, of females, 114-204 12-87 mm, of females, 17-60 mm; length of hind foot of males is 16-42 mm, of females, 17-33 mm; height of ear of males is 10-16 mm, of females, 8.8-12.8 mm. Condylobasal length of the male skull is 22.8-47.2 mm, of

Body
tail

length of males

is

mm;

length of males

is

females, 27.5-36.7

of females, 12.5-18.9

mm; mastoid width mm; interorbital

of males

is

11.0-22.4
is

mm,

width of males

5.0-10.8

971

mm,
9.0

of females, 5.5-7.8

mm;

postorbital width of males

is

6.8is

mm,

of females, 6.2-8.0

mm;

zygomatic width of males

13.8-26.0
is

mm, 12-20 mm.

of females, 13.8-19.2

mm. Length

of the os penis

647

Fig. 235. Skull of weasel, Mustela (Mustela) nivalis L.

972

Weight of males
(V.H.).

is

36-250 gm, of females, 29.5-117 gm^

Systematic Position

Weasel

is itself

an entirely specialized form; however, within the

it is,

limits of the

genus Mustela,

apparently, properly placed at the

beginning of the entire series of species.

In several respects,

less

specialized.

This

mentioned "infantile"

compared to all remaining species, shown, first of all, by the abovefeatures of its skull. As mentioned before,
it

is,

is

these infantile features are associated, to a considerable degree,

with the very small dimensions of the animal, and are particularly
distinct in the small
in its large races,

forms {rixosa group), but they are strong even which give up little in dimensions to the ermine
all to

(weasels of the boccamela-heptneri group).

The weasel stands nearest of

ing the structure of the os penis.
the details of the

the ermine (M. erminea),

although they differ in a series of essential characteristics, includIt is

probable that this

mechanism of reproductive

one of isolation where both

is
is

species are morphologically closer to each other in areas simulta-

neously inhabited by them (Tien Shan). The weasel through the ermine with M. altaica with which
in structure
it

connected

does not differ

quite a close

of the genital organs.

On

the whole,

it is

group, noteably distanced from the larger species of the genus

(V.H.).

Geographic Distribution The temperate, and in part, the Worlds; in the Old World, also
Geographic Range
in the Soviet

arctic

zone of the

New

and Old

in part the subtropical zone.

Union

This occupies essentially the entire State territory and constitutes

the greater part of the range of the species in the

Old World.

^Dimensions of body and skull

after

of L.G. Morozova-Turova; os penis dimensions after

data also from the collections of the Z[oological]

Stroganov (1962) and unpublished material Ognev (1935); weight and other

M[useum

of]

M[oscow]

U[niversity].

Minimum dimensions

of females are exaggerated.

974
In the north, the range extends to the shore of the Arctic

Ocean. The weasel is absent on the islands of the Arctic Ocean, except Bol'shoi Lyakhorsk. Occurrence of weasel on Karaginsk
Island

was not recorded;

it.

it

is

absent on the

Commanders;
is

it

is,

apparently, found on Shantar Islands, although there

information about

It
is

no positive inhabits Sakhalin (information on the abit

sence of weasel here

mistaken); of the Kuril islands,

it

is

present

only on Kunashir; the possibility that

of the southern islands
tions,
it

may be met

with in some

is

not excluded. In spite of

some
It

asseris

is

also found in the

Kyzylkum and Karakum.

its

not

excluded that the weasel does not reach the extreme north of
Taimyr, and information about
completely definite.
650

occurrence in the Pamir are not

Geographic Range outside the Soviet Union

Includes
all

Europe, including England (absent

in

Ireland), the

islands of the Mediterranean Sea (Balearic, Sardinia, Corsica,

Sicily,

Malta and Crete; apparently absent

in Cyprus), Algeria

and

Morocco (and probably Tunisia), the Azores (the weasel probably was introduced into Malta and the Azores), and Egypt (Lower). In Asia, it is encountered in Asia Minor and northern Iraq. In
Iran,
it

occupies the northern, northwestern and northeastern parts

of the country; from the northwest, the range extends quite far to
the southeast as a projection along the Zagros
exists in northern Afghanistan, Kashgaria

mountain system.

It

(known

to the east as far

as Lobnor), in Dzhungaria, in the northern part of the

Mongolian
[river]* re-

Republic including to the south the Mongolian, and, probably, Gobi

Altai

and Trans-Altai Gobi, Khangai and the Kerulen

it

gion, in northeastern China (former Manchuria), on the

Korean

Peninsula and in Japan. In China,

is

apparently absent in Inner

distribution to the south in

Mongolia and

Tibet.

The

extent of

its

the eastern parts of China is unknown. It scarcely goes far south, although one of the forms was recorded in Tonkin (North Vietnam) however as doubtful.
In the

New

World, the weasel

is

distributed in the northern

half of North

America to the northern shore of the mainland. Just west of Hudson Bay the northern border passes [westward] along a line from Chesterfield [Inlet] on the [west] coast of the Bay to
*Now
called Cherlen Gol

Sci. Ed.

975
Bathurst Inlet on the northern shore, south of Victoria Island. The

southern border is itself represented by a complicated line passing from the coast of the Pacific Ocean at Juneau through British Columbia, the southwestern corner of Alberta and northern Montana. The range includes North Dakota, a great part (except western) of South Dakota and Nebraska, Iowa, the northern part of Illinois and Indiana, Ohio, and almost all of Pennsylvania and

West

Virginia.

From

here, a small extension of the range directs

itself to the

southwest, reaching the northwestern corner of South

Carolina.

Farther east, the border passes at

first

along the Saint Law-

rence river, and then crosses the eastern extremity of Lake Huron,

going along

its

eastern shore, then along the southern shore of Lake

Erie and thence turns towards the southeast, crosses Pennsylvania

and reaches the Atlantic Ocean.

It is

absent in Newfoundland (V.H.).

Geographic Variation
Geographic variation
complicated and
in the

weasel

is

not only very great, but also

itself is

considered one of the most interesting

cases of geographic variation

among mammals. The

systematics

of the weasel then and

now

led and lead to fundamental disagree-

ments.

Even
Europe,
the

at the

beginning of the past century,

large,

it

was noted

that in

among

comparatively long-tailed weasels, there were

this

encountered very small short-tailed ones. Later

form received

name M. minuta, and began

species,

to

distinct

existing in several

be considered as a category of parts of Europe among

common weasel M. nivalis (M. vulgaris). was established that in Siberia as well as in the Far East, in Japan and in northern Europe, only the small short-tailed weasel exists. Already in the current century, it was proposed that these forms be attributed to the American "species", M. rixosa (Kuroda, 1921). Later (G. Allen, 1933), the Central and West European small weasels which were given the name M. minuta were related
populations of the larger
Later,
it

to the species rixosa. In this

way, representations concerning the

two species of weasels in the Palearctic were affirmed and each of them had its own geographical races (subspecies). Everything was complicated by the fact that, along the southern
border of the species range of the large weasel (Mediterranean), the


976
existence of a particularly large forms was recorded, which

some
(M.

authors

were

inclined

to

consider

separate

species

boccamela). This opinion, on the contrary, had been rejected

earlier, but the representation

much
in the

of species independence of the

European M. minuta was

can authors, upspecies

still

held by

some western authors

40's and even the end of the 50's (Van den Brink, 1958). Ameritill now, consider their weasel an independent M. rixosa and that the small weasels of Asia and Europe

belong to

its

subspecies.

In actuality, a long time

tists

ago

it

(Ognev, 1935, for example),

Its

that in the

was accepted by our systemawhole Palearctic only

very variable, with a

one weasel species

652

exists, geographically

complicated internal structure.

or

extreme forms (pygmaea-heptneri)

are so strongly differentiated that with limited material (territorial

quantitative)

and

insufficiently

worked

out

theoretical

prerequisites, they are actually very easily accepted as different

species. Other

European investigators gradually came

others).
It is

to

agree with

the concept of species unity (Ellermaji and Morrison-Scott, 1951;

Zimmermann, 1959 and

sels of

also doubtless that the wea-

North America are nothing more than a subspecies or group of subspecies of M. nivalis. Their specific identity with the small weasels of Asia was shown long ago (G. Allen, 1933).
Details of geographic variation, relationships between the various races, their origin and formation, etc.
clarification with large samples.
still

demand

further

At the present time,

the following

scheme of geographic

variation in weasels

may be

provisionally

accepted for our fauna. ^'

All subspecies of weasel

may be

divided into 3 groups

well-characterized morphologically and regularly localized geographically: 1) small weasels of the group pygmaea-rixosa; 2) large

weasels of the group boccamela and 3) races of intermediate type

of the group nivalis.

A. Subspecies of the group pygmaea-rixosa, small weasels. Very small weasels with very short tails and small skull of the
infantile type; in our country, they

become completely white

in

winter.

"In the description of infraspecific geographic variation of weasels, the unpublished material, kindly

made

available by L.G. Morozova-Turova,

, ,
.

was used equally

with other materials.

978
Northern European part of the country, Siberia, and Far East. USSR Finland, northern Scandinavian Peninsula, Mongolian Republic, northeastern China (former Manchuria),

Outside the

Japan, North America.

Weasels of
Carnivora.
1.

this

group are the smallest of

n.

all

forms of the

Siberian weasel, M. (M.)

pygmaea

J.

Allen, 1903 (syn.

kamtschatica, karaftensis; with respect to the weasel of the Kuril

Islands, the

name namiyei was

its

applied).

Dimensions very small.

Tail short,

length equal to hind foot or a bit longer and

constitutes about
In

13%

of body length.

summer

coat, dorsal color dark-brown, very rarely lighter,

reddish. In winter, entirely white. Hair length on sacrum, 10.3

in

mm

summer and

in

winter 14

mm.

Skull small, with weakly devel-

oped postorbital processes, short and wide postorbital constriction with weakly developed crests even in fully adult individuals. Body length of males (32), 133-M 160-1 72 mm; tail length,

12-M22.6-28 mm; hind

toid width,

foot length,
skull,

16-M21-25 mm.

22.8-M32.7-34.8 mm; mas5.0-M7.28.2 mm; postorbital width, 7.0-M8.0-8.5 mm; zygomatic width, 13.8-M16.5-19.8 mm. Weight of males (14), 37-M52.6-68 gm, of females (6), 29.5Condylobasal length of male

11.0-M 15.2-16.8 mm;

interorbital width,

M41.0-62 gm
University)'^.

(material

in

Zoological

Museum

of

Moscow

of Siberia, except southern and southeastern TransBaikaliya; northern and middle Urals, in northern Kazakhstan (a
All
653

few south of Syr-Dar'ya mouth), and Far East including Sakhalin and Kuril Islands, northern European part of the USSR westwards to Kola Peninsula and southwards to northern parts of Kirovsk and Gor'kovsk districts. A few as far as Moscow district.
Outside the USSR Finland, northern part of Scandinavian and Korean peninsulas, Mongolian Republic except eastern part and probably northeastern China (former Manchuria).
'^Data of males and females

combined

(21)*:

29.5-M49.3-68 gm. Material from

district,

Sakhalin and Kamchatka to Altai. Animals from Perm

form or approaching nivalis average, apparently, somewhat M60.8-70 gm. * Apparently includes one unsexed animal Sci. Ed.

larger:

belonging to this males (5) 50-

979
In the

are in the

European part of the Union, the most typical weasels most northern parts of the range. In the more southern

parts of the described region, weasels are locally

mixed with forms

of group nivalis or transitional populations.

It is

possible that, in actuality in the vast territory outlined, the

is

population

not systematically

homogenous and

there

may be
it

distinguished forms here differing from true pygmaea. Thus,

selves represent a distinct form

is

possible that weasels of the southern part of the Far East them(it

apparently can be given the

name mosanensis
weasels
are
is

or namiyei); the position of Sakhalin and Kuril

etc.;

not clear,

West Siberian weasels (except northern)

n.

sometimes assigned
2.

to nivalis (Stroganov, 1962), etc.

Trans-Baikal weasel, M. (M.)

punctata Domaniewski, 1926

length
consti-

(syn. kerulenica).

According
tutes about
n.
is

to

dimensions and relative

length),
it

tail

(tail

13.9% of body

corresponds with M. (M.)

pygmaea, but according to color of summer fur of upper body, somewhat lighter. In winter, it becomes completely or almost completely white. Skull somewhat wider in its rostral portion, postorbital constriction somewhat sharper, and sexual dimorphism more weakly displayed than in M. (M.) n. pygmaea. Body length of males (8), 150-M166.6-185 mm, tail length
(11),*

M20.2-28 mm; length of hind

foot,
(6),

12-Ml 8.2-23 mm'l

31.0-M32.4-35.0 mm,
6.6-M7.2-8.5

Condylobasal length of male skull

of females (8), 27. 5-M30. 4-32.0

mm;

mastoid width of males,

12.5-M14.3-16.0

mm;

interorbital width of males,

mm,

of females, 5.5-M6.3-7.5

mm;

postorbital width of males,

6.5-M7.4-8.0 mm, of females, 6.2-M6.9-8.0 mm. Weight of males (9), 36-M48.6-84 gm, of females

(7),

41-

M48.7-63 gm.'*
Southern and southeastern Trans-Baikaliya.
Outside the

USSR eastern

part of

Mongolian Republic, and

probably northeastern China (former Manchuria).

"Dimensions of animals from eastern
in

part of

Mongolian Republic. Materials

Zoological

Museum

of

Moscow

University.

from collection of Z[oological] M[useum of] M[oscow] is not shown: 30-M46.8-84 gm. Weight of female with 1 1 large embryos, 94 gm. Data on weight on material from eastern part of Mongolian Republic (Choibalsan on Kerulen). Materials of Z[oological]
'''Weight of 21 specimens

U[niversity] including individuals, the stx of which

M[useum of] M[oscow] U[niversity]. *Minimum value omitted in Russian

original

Sci.

Ed.

980

main area of distribution outside the boundaand reaches us only on the edge of its range. Lightening of body color is, apparently, connected with occupation
This form has
its

ries of the

USSR

of more arid regions.

B. Subspecies of the group boccamela, large weasels. Weasels of very large dimensions, with large skull and relatively long tail and lighter, sometimes very light color; locally, they do not turn white, or not completely white, in winter. Infantile
features, characteristic of weasels of the preceding group, are not

developed, or only weakly. Skull relatively rough, with better

developed

crests, sexual

dimorphism

is

sharply marked.

Trans-Caucasus, from western Kazakhstan to Semirech'e, and in the flat deserts of Middle Asia.
Outside the
654

USSR southern

Europe (Mediterranean), Asia

n.

Minor, Iran, Afghanistan.

3.

Trans-Caucasus weasel, M. (M.)

boccamela Bechstein, 1800.

Dimensions very
In

large, tail very long, considerably longer than

hind foot, constituting about

30%

of body length.

summer

fur,

color of upper body light brownish or chestnut,

tints. In some individuals, a brownish behind the corner of the mouth and sometimes on chest and belly. Winter fur not so pure white as in more northern forms;

with yellowish or reddish

dot

some animals in winter are piebald in color dirty white with brown patches, some do not turn white at all. Skull very large with marked constriction behind infraorbital
processes.

Body length of males (20) 209-M226.5-260 mm; tail length, 55-M72.3-85 mm; length of hind foot, 22-M39.5-42 mm.
Condylobasal length of male
females
(6), 29. 8-M3 1.2-36.0

skull,

41.0-M42.1-46.2

mm;

of

mm;

mastoid width of males, 20.0-

M20.2-22.4 mm; of females, 14. 0-M15. 1-16.8 mm; interorbital width of males, 8. 5-M9. 5-10.0 mm; of females, 6.2-M7.1-7.5 mm; postorbital width of males, 6.8-M8.8-8.4 mm; of females, 6.0M7. 1-8.0 mm; zygomatic width of males, 2 1.5-M22. 6-24.0 mm. Weight of two males, 160 and 181 gm.
Trans-Caucasus.
Outside the

USSR southern

Europe, Asia Minor, and probessential

ably western parts of Iran.

The weasel of Trans-Caucasus does not have any

differences from the South European. However,

among

the latter.

981
apparently individuals

may

often be found possessing brownish

dots on the chest and belly.

4.

Middle Asiatic, or Turkmenian weasel, M. (M.)

n.

heptneri

L. Turova, 1953.

Dimensions very
constitutes
In

large, in general corresponding with preced-

ing form. Tail very long, considerably longer than hind foot and

25-30% of body length. summer fur, color of upper

"desertish"

brownish-sandy or pale-yellowish,
mm mm

part of

body very

light,

lighter than in all

other forms of the species inhabiting our country. Fur short, sparse and coarse, in winter does not turn fully white. Length of hairs on the sacrum 5 in summer and 7 in winter. Skull large and massive with widely separated zygomatic arches. Body length of males (7), 230-M234. 1-242 mm; tail length,

55-M68.2-87 mm; length of hind

Condylobasal length of male mastoid width, 2 1.0-M2 1.3-22.0

foot,

27-M33.2-35 mm.

skull, 40. 5-M43. 1-45.5

mm;

mm;

interorbital

width, 9.8-

MlO.3-10.8 mm; postorbital width, 7.8-M8.3-9.0 mm; zygomatic width, 23.4-M24.4-26.0 mm. Weight up to 250 gm (male from southern Tadzhikistan; Z[oological] M[useum of] M[oscow] U[niversity]. Level semideserts and deserts of southern Kazakhstan and Middle Asia from Caspian Sea to Semirech'e and southern Tadzhikistan, Kopet-Dag.
Outside the
of Iran.

USSR in

Afghanistan and the northeastern part

C*. Subspecies of the group nivalis, average weasels. Weasels of moderate general dimensions, with tail of moderate length, very diverse in all characteristics of skull and body measurements. Sexual dimorphism well developed. Middle and southern regions of the European part of the country; Crimea, Cis-Caucasus and northern Caucasus, western Kazakhstan, southern, and in part, middle Urals, montane parts of Middle Asia except Kopet-Dag.
655

In all respects

as well as in distribution

subspecies

of this

group occupy an intermediate position between the small group (group A) and the large group (group B) of subspecies. 5. Middle Russian weasel, M. (M.) n. nivalis Linnaeus, 1758
(syn. gale).
Misprinted
in

Russian original

Sci. Ed.

982

Dimensions moderate, tail of moderate length about 20-21% of body length and about 1.5 times greater than length of hind
foot.

nut, rarely lighter

Color of upper body in summer fur is dark-brownish or chestyellowish-brownish. Winter fur pure white. The

hair
12.5

length

on sacrum 9.5

mm

in

summer, and

in

winter,

mm.

Body length of males (14), 163-M 186-2 13 mm; of females (9), 145-M 165-1 85 mm; tail length of males, 28-M40.5-66 mm; of females, 20-M33.6-42 mm; length of hind foot of males, 22M27.3-30 mm; of females, 19-M22.1-25 mm.
of females (9),

Condylobasal length of male skull (13), 30.5-M35. 3-40,2 mm; 29.0-M 32.0-36.8 mm; mastoid width of males,

of females, 12.8-M15.0-17.5 mm; interormm; of females, 5.0-M6.9-7.0 6.0-M7.7-9.5 bital width of males, mm; postorbital width of males, 5.0-M8. 2-9.3 mm; of females, 5.3-M7. 1-9.0 mm; zygomatic width of males, 11.0-M17.9-22.0

15.0-M17. 3-19.6

mm;

mm;

of females, 15. 3-M16. 2-18.5 mm. Weight of males 60-100 gm (probably somewhat more).

Middle regions of European part of the USSR from Pribaltic to middle and southern Urals, northward approximately to the latitude the of Leningrad and Perm, and south to Kursk and Voronezh districts. Outside the USSR northern parts of Europe, except Finland

and northern part of Scandinavian Peninsula. This form does not possess very definite characteristics and itself represents, on the whole, one of the stages in the transition from the small Siberian and North European form pygmaea to the larger form vulgaris and further to the large southern weasels. The area of intermixing and transition of individuals both toward one (pygmaea) and the other (vulgaris) tendencies, i.e. in the north and
in the south, is vast.

As was shown,

individual animals of the type

pygmaea

are found to the north of Gorki and Kirov

districts, and, at the

Moscow
trict are

and even same time, the weasels of Perm disto,

already close in dimensions

or belong to, nivalis.

Due

form is very great, and equally forms, the extreme small "normal" large relatively with the so of the subspecies and of the range variants are found in nearly all even to Middle and Western Europe (form minuta). Equally with this, individual animals and separate small populations of relatively large weasels are found in the range.
to this, individual variation of this

983

The
form
is

possibility of accurately diagnosing

and designating

this

made more

difficult

by lack of

clarity

concerning represento the

tation

of the typical nivalis lineage, which, probably, itself

represents a transition from the northern

pygmaea

Middle

European vulgaris. The form nivalis is conditionally accepted here. 6. Middle European weasel, M. (M.) n. vulgaris Erxleben, 1777
(syn. nikolskii, dinniki, caucasica, trettaui).

Dimensions somewhat larger than

relatively longer,
its

in

M. (M.)

n.

nivalis. Tail

length constitutes on average about

27%

of

body length and approximately twice as long as the hind foot.

Color of upper body in summer fur varying from lightbrownish to dark-chestnut. Winter fur white and only in extreme south of range is whitening incomplete. Hair length on sacrum
7.5-8.5
in

mm

in

summer and

in

winter 10.5-13.5

mm.

Skull larger

M. (M.)

n. nivalis,

and zygomatic arches relatively widely sepa-

rated.

656

Body length of males (26), 173-M2 12.6-243 mm; of females (6), 155-M181. 3-212 mm; tail length of males, 45-M57.9-75.5 mm; of females, 45-M54.1-64 mm; length of hind foot of males, 21-M30.8-38 mm; of females, 29-M25.0-31 mm.
of females (6), 25. 8-M33. 2-36.0

Condylobasal length of skull of males, 31.5-M38. 8-43.0 mm; mm; mastoid width of males, 13.5-M18.4-20.5 mm; of females, 14.0-M15. 1-16.8 mm; interor-

bital

width of males, 6.5-M8.5-10.2

mm;

of females,

6.2-M

7.1-7.5

mm;

postorbital width of males,

females, 6.0-M7.1-8.0

mm;

5.0-M8.4-10.2 mm; of zygomatic width of males, 17.0-

M20.0-22.0 mm.
latitude of southern

Southern regions of the European part of the country from the Voronezh and Kursk districts, Crimea, Cisat least

Caucasus, northern slope of the Main Caucasus, eastward to the Volga (eastern border not precisely known).
Outside the
enees.
In

USSR Europe
its

southward

to the

Alps and Pyr-

some

parts of

range

in the south, the

described form has

features significantly similar to large weasels of the type boccamela.

This reveals

itself particularly

partially explained

in the northern Caucasus, and is by mixing of the population of large Trans-

Caucasian weasels penetrating northward, hybridizing with them, etc. An analogous phenomenon is observed in roe deer (see Vol.
I

of the present series).

984

The form vulgaris

the north, the

in

the accepted sense itself represents a

boccamela. In form vulgaris insensibly flows together with the form nivalis over a broad zone and distinguishing them is only possible through series. The acceptance of two forms is, to a certransition to the large southern weasels of the type
tain degree, conditional.

The majority of western European authors accept for "Germany" the form trettaui. This is nothing more than a synonym of
the

form vulgaris and

is

connected with nothing other than the

different nomenclatural interpretations of the

name

vulgaris,

believed to be a simple

which is unfounded. In general, for a correct understanding of the form nivalis,

nivalis, a fact

synonym of

a solid revision of the weasels of the Scandinavian Peninsula

is

necessary.
n.

Some

authors consider the Scandinavian weasel

M. (M.)

pygmaea

as identical to the

Middle European "minuta" and an

n. nivalis

extreme variant ("minus- variant") of /. (M.) 1957), with which it is difficult to agree.
7.

(Reichstein,

Montane Turkestan weasel, M. (M.)

in

n.

pallida Barrett-

Hamilton, 1900.

Dimensions somewhat smaller than

of moderate length, constituting about

M. (M.)

n.

vulgaris,
Tail

approximately correspond to dimensions of M. (M.)

n. nivalis.

24%

of body length.

Color of summer fur light-brownish. Winter fur white. Skull of moderate size, relatively narrow in mastoid part, postorbital constriction relatively short

and narrow.
length,

Body length of males (17), 180-M 192-2 15 mm, tail 37-M47-60 mm; length of hind foot, 25-M27-31 mm.

Condylobasal length of the male skull, 31.8-M35. 4-40.0 mm; mastoid width, 15.0-M16. 6-19.0 mm; interorbital width, 6.5-M7.7-

mm; postorbital width, 6.5-M7.5-8.5 mm; zygomatic width, 18.0-M19.7-22.0 mm. Weight of males (1), 113 gm, of females (3), 75-108 gm. *Outside the USSR found in the Chinese parts of the same mountain systems, and perhaps in extreme eastern parts of Hindu8.4

kush (Afghanistan).

The scheme presented focuses on

in

the
in

main features and trends

our country.
It

geographic variation of the species

is

cannot be

considered sufficiently studied, and

"Description of range within

not generally accepted (see

montane
Sci. Ed.

parts of

USSR omitted in Russian original; this includes Turkmenia, Uzbekistan, Tadzhikistan, Kazakhstan, and Kirghizia

985

Ognev, 1935; Novikov, 1956; Stroganov, 1962). The question of weasels in the European part of the country and Caucasus is particularly complicated.

657

Pyrenees Peninsula and 1906 M. (M.) galinthias Bate, Balearic Crete; M. russeliana Thomas, 1911 Sichuan, southern China; M. (M.) namiyei Kuroda, 1921 Japanese M. (M.) (M.) M. (M.) mosanensis Mori, 1927 Korean Peninsula; tonkinensis Bjorkegren, 1942 northern and southern Vietnam race belongs another probable 10) M. (M.) alleghaniensis Rhoads, 1901'^ southeastern of the range
siberica Barrett-Hamilton, 1900
Islands; 5)
n.

For the parts of the range lying outside the USSR, the followM. (M.) n. subpalmata Hemprich Egypt (Lower); 2) M. (M.) n. numidica Pucheran, et Ehrenberg 1855 Morocco, Algeria, Malta, Azores Islands? Corsica; 3) M. Kashgaria'^; 4) M. (M.) n. (M.) n. stoliczkana Blanford, 1877
ing forms are usually accepted: 1)

[Iberian]

6)

n.

7)

n.

Islands; 8)

n.

9)

n.

(it

is

that this

to

species);
part

n.

America (Michigan, Pennsylvania, 'Virginia, North Carolina, Illinois, Wisconsin); 11) M. (M.) n. campestris Jackson, southwestern angle of the range in America (South Dakota, Iowa, Nebraska); 12) M. (M.) n. eskimo Stone, 1900 Alaska; 13) M. (M.) n. rixosa Bangs, 1896 remaining major part of the range in America (Mackenzie, Labrador, Quebec, Minnesota, North
in

Ohio, 1913

Dakota, Montana, Saskatchewan, Alberta, British Columbia) (V.H.).

Biology
Population. Within the boundaries of
its range, the weasel is everywhere quite common, but unevenly distributed, animal. This small carnivore serves as a good indicator of the abundance of mouse-like rodents almost its sole food. At the same time,

wherever ermine are abundant, weasels are few and vice versa (Ognev, 1931). This applies to the distribution of both numbers
''Information about the existence of this form in the USSR in Semirech'e at Dzharkent (Ellermann and Morrison-Scott, 1951) with reference to Ognev (apparently 1935), is based on misunderstanding. In describing this form, S.I. Ognev nowhere spoke about its presence within the boundary of the USSR and refers to Yarkend (Kashgaria) and not Dzharkent* (Semirech'e).

*Now
'*The

Panfilov

Sci. Ed.
this

American authors consider separate species M. (M.) rixosa.

and the following form as belonging

to a

986
inside the range and density in separate stations at the limits of the
area. In the

upper Pechora, one weasel track

is

found for 10-18

tracks of ermine; on the contrary, in the southwestern part of Kalinin

district,

3-4 tracks of weasel for

1

track of ermine, and in Zhiguli,

7 weasel tracks for

ermine.
lies

The area of weasel abundance

more southward than

is

that of

ermine, and numbers of ermine in southern regions

bilities

usually less

than weasel numbers. This partly depends on differences in possi-

of capture of mouse-like rodents between these two spe-

cies. In the

European

part of the

USSR, weasels

are found in small

mainly live in and around human settlements, and are rare in the forests. In the intermediate zone, it decidedly predominates over the ermine. This is still more obvious in the forest-steppe belt and the chernozem steppes. In the Caucasus, the weasel completely predominates. In
in the taiga forest zone. In this zone, they fields

numbers

Siberia,

it is

Siberia,

it is

more numerous in the southern parts. In northeastern rare. It is more often encountered in southern Transnumbers, moreover not oneits

Baikal and Primore Territory.

Due
time,
its

to significant fluctuations in

occurrence in different parts of

range can only be

658

compared under similarly favorable environmental conditions. Table 62 gives the frequency of weasel tracks (per 10 km) for the mixed forest zone and for broad-leaf forests. Extreme magnitudes apply to different habitats in one season. In both cases, frequency of tracks may change 8-10-fold, but in all cases, in broad-leaf forests, it is 3-4 times more than in mixed forests. Habitat. The weasel is not finicky, and can accommodate itself easily to any conditions. Wherever there are many mouse-like
rodents, one

may

find the weasel.

is

It is

established in tundra, does

not avoid taiga and

common

in

the forest-steppe zone and

chernozem
tains. It

steppes.

It

lives in grassy steppes, semideserts

to

does not avoid proximity

humans,

settling

and mounsometimes in

Table 62. Frequency of weasel tracks in the broad-leaf [Zhiguli] and

mixed forests [Kalinin] (along 10

Year of high numbers

km

route)

Year of low numbers

Region
Extreme
Southwestern
Kalinin
Zhiguli
part,

Quantity

Average
9.7
36.2

Extreme
0.2-8.6
0-13.3

Average
0.9
2.7

0.7-12.8
9.7-122.8

district

^ ^

^^ >

^
^
^N

~ ^

ii
^^ =1 a _" ^ U ^^
tt

^
><-

__"

u
ce

(U

00

2 (
S 3

5 z

g ^
_;

t:
'

Z <

^ 2

:g

a u
<e

a-

T3

987
structures within a

town

(outskirts of

Moscow). Although encounis

tered almost everywhere, the weasel

known

to prefer certain
is

places. Thus, in the middle forest belt, the weasel

most often

met with

in

openings, clearings, forest edges, plantations, roadside

It is

ditches, etc.

often in riparian vegetation of small rivulets, in

meadows with
sel
is

haystacks, in mezha*, threshing-floors**, in store-

houses, barns, and other buildings in villages. In Siberia, the wea-

mainly
the

more often encountered in steppe than in taiga, where it restricts itself to meadows, old burns and in tall grass. In chernozem and steppe belts, it inhabits steppes, fields, pasvegetable gardens, stonepiles, woodpiles, thresh-

tures, gardens,

ing-floor**, hay/grain stacks, fences, inhabited and uninhabited

structures;

der fields,

mountainous places among talus slopes, bouland semideserts, the weasel is closely associated with the places where colonies of social rodents are
and
in
etc. In deserts

found, mainly gerbils (particularly great gerbil [Rhombomys opimus])

and simply

lives

among

these colonies or in

them (V.G. Heptner).

Food. The weasel basically feeds on mouse-like rodents: voles, mice, hamsters, gerbils and others. Food composition is variable,

and depends upon which of them predominates in the given locality. The weasel does not always decide to attack adult hamsters and rats. The weasel usually bites small voles in the occipital
region of the skull. This
is

the only

way
it

it

kills its prey. It

can bite
it

through [the skull of] a young water vole only with difficulty;

can deal with pikas and gerbils, but

cannot overcome young

Norway

rats

and sousliks.
is
it

Species composition of prey

the weasel

also determined

by habitat of
in forest than

in the

same

locality,

behaves different

near villages and in the fields.

is

characteristic feature of the weasel

the fact that in

its

food, shrews are

more numerous than

in other

carnivores (Formozov, 1948); in different years in Tatariya, their

frequency reaches
In

25%

(Grigor'ev and Teplov, 1939).

of the weasel consists of gray

Murmansk

district, the diet

[Microtus]

[Lemmus]

and red-backed [Clethrionomys] voles, Norway and wood [Myopus] lemmings, and rarely fish

*Strip of uncultivated land between

**Two Russian words,

Sci.

tok and

two fields Sci. Ed. gumno, are both translated

as "threshing-floor"

Ed.

988
(Nasimovich, 1948). In Trans-Baikal, the weasel feeds on narrowskulled voles [M. gregalis] (50%), Asiatic

wood mice [Apodemus

peninsulae] (25%), bank voles [C. glareolus] (12.5%) and small

birds (12.5%) (Fetisov, 1942). In the steppes of the northern parts

of the Mongolian Republic, in years with an abundance of Brandt's

vole [Lasiopodomys brandti], the weasel feeds mainly on
as
it

as well

659

on the clawed Mongolian gerbil [Meriones unguiculatus] (Kucheruk, 1948); in the deserts of Pri-Balkhash on the great gerbil [Rhombomys] (Sludskii, 1953). In the forest-steppe of western Siberia, small voles, field mice, Dzhungarian hamsters, [Phodopus sungorus], water voles enter the weasel diet, and in small amounts, water voles and fish, and even carrion is sometimes met with (9.1%; Zverev, 1931). In Tatariya, the common vole predominates (23.1%) in the weasel diet, while other voles, including water, and field mice, are utilized in lesser quantity. The average percentage of occurrence of brown-toothed shrews [Sorex] equals 17% (Grigor'ev and Teplov, 1939). In the middle [forest] belt, the weasel prefers the common vole and field mice. In captivity, water shrews [Neomys] are very unwillingly utilized (V.V.

Kucheruk). Frogs,

fish,

small birds and bird eggs are very rare in

terrestrial

weasel food. In Ussuri Territory, the weasel often eats

the seashore (Yu.A. Salmin and V.O. Shamykin).

molluscs and feeds on a number of marine organisms cast up on

In the literature are

known

cases of successful attack by

weasels on larger prey larger than the weasel itself up to capercaillie, hazelhen and hare. However, such cases occur extremely rarely. In the stomachs of weasels are the remains,
at

one

time of only one mouse-like rodent; in

than 10

it

usually occur no

more

gm
the

of food (Griror'ev and Teplov, 1939), a weight con-

stituting about 1/10 of the weasel's

body weight. The

daily [food]

norm of

about 30body weight (V.V. Kucheruk). However, weasel kills significantly more mouse-like rodents and shrews than needed for
little

animal

is

equal to about 35 gm,

i.e.

35%

of

its

in abundance, it eats only a small sometimes only the brain. According to the method of obtaining its food, the large Caucasian weasels undoubtedly belong to the ermine type, since it is impossible for them to penetrate into the holes of small rodents because of their

food.

When

its

prey

is

found

part of captured animal,

large dimensions.

Home

range: The dimensions of the weasel's

home range

are

determined by the abundance of the prey within

its limits,

and the

989

659

Fig. 238.

Turkmenian weasel, Mustela (Mustela)

nivalis heptneri L. Tur*., looking

in

out from a burrow of a great gerbil,

Rhombomys opimus. Kyzyl-Dzhar

Badkhyz

preserve, southern Turkmeniya. 28 April 1963. Photograph by Yu.K. Gorelov.

daily range depend also on weather conditions.

The home range

is

usually distributed
ity.

among

a series of adjacent areas of daily activfor the

The home ranges of males and females

most part occur

contiguously. In the forest-steppe of western Siberia, the area of

660 daily activity

Murmansk

sometimes comprises 0.5-0.6 ha (Zverev, 1931). In dimensions of the home range of weasels in winter did not exceed 10 hectares in many cases. The length of the daily route in this case ranged from 1.5-2.5 km (Nasimovich,
district, the

1949).

Burrows and

shelters. In winter, the weasel has

no permanent

burrows or other shelters. In Murmansk district, when at rest, it always lies beneath the protection of the snow in the roots of a spruce tree, among stones, and under fallen branches (Nasimovich,

1948). In the middle [forest] zone, these occur as woodpiles, heaps

of brushwood or fallen branches, straw stacks or piles of hay in

* In Russian original, misspelled

""., but author of name, heptneri is

Morosova

Turova; see

p.

Sci. Ed.

990

meadows; and finally, quite often, the subnival nest of a vole which the weasel has eaten (Tikhvinskii, 1936). Usually within the limits of the home range, the weasel has some shelters which it
regularly visits.

on brood burrows. There are indications that nests of weasel with broods were sometimes found under old rotten stumps or among sheaves and straw remains of the previous year. The weasel often settles with its brood in nesting chambers of vole burrows. Daily activity and behavior. The weasel has no defined regime of daily activity. It is, for the most part, active at twilight and at
There
is little

reliable data

night, but

it

or

its

fresh tracks

may be met

during daytime, espe-

cially during a snowfall or

snowstorm.

In case of food abundance, the area of daily activity is densely

covered with

its

tracks in a very small area.

At

night,

it

does not

pass by any fallen log, bush, stump, hole, or heap of branches

without inspection.

When

its

prey

is

scarce, the tracks of the weasel

660

Fig. 239.

Turkmenian weasel, hunting

reserve, southern Turkmeniya.

in a

colony of great gerbil. Kyzyl-Dzhar

is

in

Badkhyz

steppe agarna

between the teeth of the

weasel. 26 April 1963. Photograph by Yu.K. Gorelov.


991

extend along an almost straight line for hundreds of steps to another

more prey-rich

area.

The weasel usually hides

for a long
it

period in subnival burrows of voles and shrews. Very often,

uses the underground passages of moles. In severe frost or crusted

snow, the weasel almost never appears on the snow surface.

always catches
its

It

almost

prey under the snow and rarely on

its

surface.

661

The weasel moves exclusively by jumps, with the distance between tracks of the fore and hind feet from 18-35 cm, depending on the leap and the size of the animal. In Middle Russia, length
18-20 cm, the feet stand close in the leap. Given the mice and voles in excess of its daily needs, the weasel often makes stores, sometimes very significant, and usually in one place, where from 1 to 19 voles and mice may be found. In 13 cases, their average number was 8 (V.V. Kucheruk). Reproduction. Reliable data on the time of mating and the duration of the rut period are lacking. There are also no data on the duration of pregnancy, as well as on its nature. To all appearof the leap
is

possibility of killing

ances,

all

stages in the reproductive cycle of the weasel, except

possibly the duration of pregnancy (from the

to parturition) are inconstant and variable.

moment
of
all

of placentation

They

are determined

by

factors of the external environment,

and

first

by food abun-

dance.
In Silesia, pregnant females

were found

in January, April,
in the

August, October and November (Pohle, 1909);

in April,

Caucasus

and October (Satunin, 1915). In southeastern TransBaikal, pregnant females as well as those which had just given birth to young were simultaneously found at the end of June and the end of July (P.B. Yurgenson). In Kostromsk district, a female with five fetuses was caught in spring (Formozov, 1948). In Tatariya, two nursing females were caught on 7 April and 1 May, and two pregnant ones on 2 and 5 May. The nursing females had

May

4-6 placental scars and in the pregnant ones, the same number of embryos were found (V.A. Popov). On 14 April, a nursing female was caught (V.V. Kucheruk). In Kazakhstan, pregnant and post-partum females were found in May (Sludskii, 1953), and in northern Kirghizia, young weasels had almost reached adult size by the end of June (Kuznetsov, 1948). According to data of the Kazan Zoo (1929), the minimal duration of pregnancy was 7-8
weeks. Thus, pregnant females were found in various parts of the
range in
all

months of the year, but more frequently

in spring, in

992
April.

Young were caught

in

March, May, June, July, November

and December, and in the German Democratic Republic and the Federal Republic of Germany, also in September and January.

4 to 7 young occur, rarely were 3, 8 or 9 observed. There is a written record of 10 young found in a burrow (Novikov, 1959). The number of males in litter is usually

Most

often, litters of

more than

that of females.

The number of young

in the litter de-

pends on abundance of food in the given season. In years with abundant mouse-like rodents in the Mongolian Republic, there were
significantly

more embryos

(11,

11,

15,

16) than

were found

in

years of their depression (5, 7).

The maximum known number of

embryos
less,

is

19 (V.V. Kucheruk).

Growth, development, and molt. Weasels are born blind, helpcovered with scarce whitish underfur. They stay with the mother for a long time. The brood disperses at the end of summer or in autumn, when the young are already comparable in size to the
mother.

Molting proceeds twice per year

in spring

and

in

autumn.
All

Enemies, diseases, parasites, mortality,

first

and competitors.

carnivorous animals and birds feeding on mouse-like rodents, and

of
all

the ermine, are competitors of the weasel.

Many

of
it.

these are, at the

same time, enemies of

in

the weasel, destroying

Remains of eaten weasels occur

and others. Such cases are
mation that the weasel
is

excrement and debris of red fox, sable, steppe and forest polecats, ermine, eagle-owl, buzzard
rare, but nevertheless

occur in places.
is

Diseases of weasels are almost unstudied. There

tularemia infection was isolated from
resistant to tularemia (Olsuf'ev
is

some
is

infor-

susceptible to rodent plague; a culture of

it,

but the weasel

quite

and Dunaeva, 1951). Skryabingilosis

in

widespread among weasels, as also

ermine.

Rates of mortality and longevity have not been studied. Undoubtedly, in years of sharp decrease in numbers of mouse-like
rodents,

many weasels

die

from hunger.
in

Population dynamics. Fluctuations

numbers of weasels

is

well-demonstrated. They are very closely connected with the abun662 of the

dance of food mouse-like rodents not of only one species, but whole number of small mammals (voles, mice, shrews; Formozov, 1948). In a series of cases, as in the majority of the
carnivorous animals,
its

population apparently increases in the year

following one of food abundance (Migulin, 1941; Formozov, 1948

993

and others). But this does not always occur. Thus, in southeastern Trans-Baikal, in 1939-1950, the weasel population increased simultaneously with growth in the abundance of mouse-like rodents. The same was observed in the southwestern part of Kalinin district in 1948-1950 (P.B. Yurgenson). This may be explained by
the unstable and prolonged period of reproduction of the animal.
In southeastern Trans-Baikal, within a nine-year period, the

of weasels rose three times (P.B. Yurgenson). In

trict,

number Kostromsk dis-

over a 10-summer period,

it

was observed

that the

number of

weasels regularly increased every other year (Formozov, 1948); after a year of abundance, numbers fell sharply the following year.

Rapid increase
fluctuations

in

numbers

is

explained by high

fertility,

and sharp

by narrow feeding specialization (voles and shrews).

number Mongo-

In southeastern Trans-Baikal, within a nine-year period the

of weasels changed by 2-3-fold (P.B. Yurgenson). In the

lian Republic,

crease in

between 1944-45, there was noted a nine-fold innumber of weasels (Kucheruk, 1948). In May 1944, in one hectare of the test areas, an average of 0.14 weasel was ob1.27 weasels. In April 1945, 0.66; in September no weasels were caught in any of the test areas. In Kostromsk district, during one year the weasel population increased and decreased 10-fold (Formozov, 1948). During the period from 1930 to
tained; in June

1950, the weasel population in the southwestern part of Kalinin

district

changed 34-fold (1934/35 and 1947/48; P.B. Yurgenson), following the end of the war, occurrences were very low 0.28-0.58/10 km route. In the upper Pechora, a 25 [fold] short-term fluctuation of the weasel population was observed
and
in the year

(Teplov, 1951).

Field characteristics.
cases, the weasel

It is

very easy to confuse the track of a

large male weasel with those of a small female ermine. In such

is

distinguished by

its

considerably smaller area

trail

of daily activity, the shorter daily length of the

and also the

very uneven restless

movement

in short

(5-10 m) wave-like zigzags,

while in ermine, the angles of turning are more acute (Nasimovich,

1949). In walking, ermine sometimes "trot", which in weasels has

never been observed. The weasel places

other than ermine, and in short

its

feet closer to

each

jumps

the tracks of "landing"and

"bounding up" often join

in

weasels in the eastern Altai (Dul'keit, 1956)

one chain. The length of leaps of is 15-25 cm, the

994

Fig. 240. Paired foot impressions of

nivalis L.,

male Middle Russian weasel, Mustela (Mustela) on snow and the pattern of leaps during quick movement of the animal. Fomkino, Moscow district. 8 December 1963. Sketch by A.N. Formozov, nat. size.

A)

Fig. 241.

legs

on the

group of tracks of the Middle Russian weasel, jumping easily on four first snow. Fomkino, Moscow district. 15 November 1951. Sketch by A.N. Formozov, about 2/3 nat. size.

995
663

diameter of the individual track

is

1.0-1.6

cm
is

(in

ermine, 1.8-2.5).

The weasel sinks load on 1 cm^ of

into the

snow

for about 1.5-2.5 cm.

The weight
(P.Yu.).

the surface of the track

7-8

gm

Practical Significance

The significance of the weasel

like rodents is very great. This fact that, in years of

in the destruction of
is,

harmful mouse-

in particular, strengthened
kills tens

abundance,

it

by the and hundreds of times

more rodents than it can eat. According to some data, a weasel can kill up to 2000-3000 rodents in a year. In "eating places" of the weasel, up to 456 half-eaten voles and mice were found among the stacks of straw on the threshing floor. The weasel hunts rodents
not only in fields and forests, but also in the villages

in

hay piles

of straw stacks, storehouses and store-rooms. Almost everywhere,

where mice and vole can penetrate, the weasel can

In the fur trade, the role of weasel
is

also.
It

negligible.

is

only

accidentally captured in traps set for ermine, Siberian weasel, etc.

The weasel

fairly often gets into traps

placed for moles. In a year

of abundant weasels, one weasel was caught against 25-40 moles

(Formozov, 1948).

As

a destroyer of harmful rodents, the weasel

its

must be pro-

tected and preserved throughout the year, and

hunting must be
in fur prepa-

prohibited everywhere.

Its

trade should be everywhere prohibited,

and the accidentally captured weasel must not be used

ration to avoid encouraging
its

hunting (P.Yu.).

ERMINE
Mustela (Mustela) erminea Linnaeus, 1758
1758. Mustela erminea. Linnaeus. Syst. Naturae, ed. X,
1,

p.

46.

Sweden.
1792. Mustela erminea aestiva. Kerr. Animal Kingdom,
p.

181.

Germany.
1816. Mustela herminea. Oken. Lehrb. Naturg.
3, 2, p.

1026. Re-

naming of erminea Linnaeus.

1857. Putorius kaneii. Baird.

Mammals North Amer.,

p.

172.

Arikam

Island in Bering Sea. Apparently,

Arakamchechen

Island (V.H.).


996
1895. Putorius ermineus ferghanae. Thomas. Ann.
15, p. 452.

Mag. Nat.

Hist.,

1896. 1912. 1913. 1914. 1922. 1922. 1923. 1928.

1928.

Kara-Karyk mountains, Fergana. (?V.G.). Putorius arcticus. Merriam. North Amer. Fauna, 11, p. 15. Point Barrow, Alaska. Mustela lymani. Hollister. Smiths. Misc. Coll., 60, 14, p. 5. Tapucha*, on the Chuisk Trail, southern Altai. Mustela nippon. Cabrera. Biol. Soc. Espan., 13, p. 392. Sinano, on Hondo (Honshu) Island, Japan. Mustela kanei. G. Allen. Proc. New Engl. Zool. Club, 5, p. 58. Nizhne-Kolymsk. Putorius erminea var. kamstschatica. Dybowski. Arch. Tow. Nauk. Lwow, 1, p. 349. Nom. nudum. Putorius erminea var. sibirica. Dybowski. Ibidem, p. 349. Nom. nudum. Nee Pallas, 1773. Arctogale erminea tobolica. Ognev. Biologich. izvestiya, I, Former Tobol'sk governance. p. 112. Mustela erminea transbaikalica. Ognev. Memuary zool. otd. Obshch. lyubit. estestvozn., antrop. etnogr., 2. Sosnovka, Barguzinsk Preserve. Mustela erminea orientalis. Ognev. Ibidem, p. 15. Pokhodskoe S. [village] on the Kolyma [river], 6904' N.

lat.,

160''55' E. long.

1929. Mustela erminea baturini. Ognev. Izv. Tikhookeanskoi

^^'^

nauchno-prom. stantsii, 2, No. 5, p. 9. Great Shantar Island in Okhotsk Sea, lower Anaur river. 1931. Mustela erminea birulai. Martino. Ezhegodn. Zool. muzeya

AN

SSSR,

31,

1930,

p.

208.

Aktyubinsk.

Nomen
p.

praeoccupatum

Kolonokus alpinus birulai Ogn., 1928.

11.

1932. Mustela erminea ognevi. Jurgenson. Zool. Anz., 98, Mouth of Taz river, western Siberia. 1935. Mustela

erminea

schnitnikovi.
3, p.

prilezhashchikh stran,

37.

Ognev. Zveri SSSR Former Kopal'sk Co.,

Semirechensk governance.
1936. Mustela erminea karaginensis. Jurgenson. Byull. Mosk.

Obshch.

ispyt. prirody, 45,

No.

3, p.

240. Karaginsk Island

near eastern shore of Kamchatka.

1938. Mustela erminea naumovi. Jurgenson. Tr. Altaiskovo gos.

zapovednik,
66**

1,

p.

124. Voevoli Lake, source of Khatanga,

N.

lat.

*In Russian original, misspelled

Topuchaya

Sci. Ed.

997
1941. (1939). Mustela erminea teberdina. Kornejev. Acta Mus.
Zool. Kijev,
1, p.

174. Teberdinsk Preserve, Caucasus.

S,c.,

1944. Mustela erminea digna Hall. Proc. Calif. Acad.

23,

No. 37,

p.

559. Kamchatka.

1951. Mustela erminea martinoi.

Checklist Palaearct. Indian

EUerman Mamm., p.

et

Morrison-Scott.

256. Substitute for

M.

e.

birulai Martino.
1,

1962. Mustela erminea balkarica. Baziev. Zool. Zhurn., 41, No.

p. 123.

Chegem

river gorge, northern slope of Great

Cauca-

sus (V.H.).

Diagnosis
Color monotone white or dark-brownish tone above and white below, with dorsal and ventral colors sharply demarcated. Distal
half of half to
tail

black. Tail length with terminal hairs comprises about

a third of

more than

body

length. Skull moderately elon-

gated

distance between the mastoid processes approximately equal

condylobasal length, but somewhat more elongated than
less than inter-

to half of in weasel.

orbital

Width of skull above canines noteably space. Dimensions are small (V.H.).
Description

In

its

general proportions, manner of posture and movement,

is

the ermine

entirely similar to the weasel

it.

and represents a someis

what enlarged copy of

a third of

However, the
its

tail

relatively longer,

never being equal to the length of the hind foot, always exceeding

body

length, and

length together with terminal hairs

length.
silky, but quite closely-lying

constitutes about half of

body

Winter fur very dense and

summer fur rougher, around Moscow, length of

short;

shorter and sparse.

Among

and ermine

17
I

mm

(M15.75), and

in

winter to 15
II

mm

directional hairs in winter is up to summer, M13.9; guard hairs of category (M13.7), and in summer, M12.1; guard hairs
in

in

category

in winter to

14

mm

(M12.2), in

guard hairs

in

category

III in

winter to 13

M10.5; guard hairs in category IV to 12 M9.9; length of underfur in winter M9.1 mm, in total number of hairs in 0.25 cm^ in winter comprises 4808, of

mm (Ml 1.9), in mm (Ml 1.5), in

summer Ml 1.3; summer summer summer 8.1. The

998

which two* are directional hairs while underfur hairs number 1 for every 27 underhairs; in summer, on 0.25 cm^ there 4 637 are 3,636 hairs, of which there are 5 directional hairs and under-

number 3,539 1 for every 36.5 underhairs. Thickness of ermine in summer decreases by 25% while in the majority of species, it decreases by about two times (Pavlova, 1959). Difference in character of winter and summer fur in southernmost forms is less than in northern. The tail is covered by short closelyhairs

fur of

lying hairs, not fluffy, and relatively fine and even throughout

whole length of the

665
ible. In

tail.

Soles of feet furred and in winter fur, their pads are not vis-

summer, pelage
color
is

is

less

dense and pads are bared.

In

summer

two-toned

on the dorsal side of the body,

the top of the head and the sides are brownish in color of various
densities and tones; the underside, the inner sides of the feet and

the lower part of the neck and head are white with greater or lesser

yellowish or lemon-yellow film; in extreme cases, the belly

bright yellow.

is

quite

The

distal half

of the

tail is

black in winter and in

Fig.

242.

Ermine, Mustela (Mustela) erminea A.N. Komarov.

L.,

in

summer

fur.

Sketch by

*Sic; this

number may be

in

error

Sci.

Ed.

999

summer, the upper and lower parts of the basal half correspond with the color of the back. The dorsal surface of the feet of both fore and hind legs are white or yellowish-white. Individual variation in color is not great. There are no age or sexual differences in color. Geographic variation in color of the

summer

fur

is

noticeably expressed, but

is

not great; color changes

from dark-tawny and dark-brown

noted, seasonal variation
a
is

to quite light
is

straw-brown. Geonot expressed.

all

graphic variation in color of the winter fur

As
in

great and expressed in

races.

Only

few of the southernmost populations or individual animals

these populations do not turn white or partially white in winter.

relatively
Its

The ermine skull is very similar to that of weasel, but more elongated, with a less broad and swollen braincase.
is

anterior region (in the region of the frontal bones)

not so

wide and voluminous and is more elongated. The postorbital constriction is sharper and somewhat longer. The facial portion of the skull width above the skull is elongated and relatively narrow

canines

is

notably less than width of the interorbital expanse. The

thin.

zygomatic arches are weak and

Zygomatic width approximastoid region. The their infraorbital foramina are rounded and relatively large
mately corresponds with skull width
transverse diameter
in the

is

considerably larger than the longitudinal

diameter of the canine alveolus.

The upper
667

profile of the skull in the region posterior to the

is

supraorbital processes
region, there
is

flattened,

and therefore

in the interorbital

an evident elevation, from which the profile of the

The prominences, crests, etc. weakly defined, but relatively somewhat stronger than in weasel the better defined is the sagittal crest found in the postorbital (frontal) region, and the occipital crest is well defined. The auditory capsules are narrow and high, their inner edges parallel to each other. The distance between them is less than in weasel, being less wide than the hard palate at the level of the anterior edge of the sphenopalatine notch. Teeth are small, but relatively
facial part quite abruptly descends.

are

somewhat stronger than

in weasel. Carnassial teeth are well devel-

oped, the upper canines longer and thinner and the lower canines

somewhat massive.

On
crest,

the whole, because of

its

its

relative elongation, less swollen

braincase and

flatness, elongation of facial part,

etc.,

development of

prominences,

the ermine skull has a less "infantile"

1000

666

Fig. 243. Skull of ermine,

Mustela (Mustela) erminea L.

1001
character than the weasel skull. This applies both to the larger and
the smaller races of the species.

Age changes
young ermine

in the

ermine skull are quite considerable. '' The

skull (first winter of life), besides

is

somewhat smaller

general dimensions,

distinguished by a relatively short facial part,

rounded and somewhat swollen braincase, weak supraorbital processes and shorter postorbital constriction. The pace of age changes in the
skull
is

quite rapid, and in the second winter of

its life,

the animal has

its

a skull which cannot be differentiated from that of adults in

main
in its
its

features (for age changes, see also description of the os penis).

Sexual differences are considerable and are manifested

somewhat
narrower
against

different proportions of the female skull besides

is

generally smaller dimensions. In females, the skull

somewhat

in the nasal part (about 23.3% of condylobasal length 24.1% on average in males), mastoid width is somewhat less (about 54% against 55.7% on average in males), in females, the facial part is shorter (about 45.5% against 48.7% in males), the cranium is relatively somewhat longer (59.2% against 55.3%). Crests in

females are usually not defined, occurring only

in old individu-

and even so are relatively less than in males (Morozova-Turova, 1961). The female skull is not so massive and is absolutely and on average lighter. The weight of male skulls (120) without lower jaws from Kamsko-Bel'sk flood lands is 1.9-M2.47-2.7 gm; that of females (70) is 1.3-M 1.54-1. 9 gm. The comparative weight of
als,

male skull is 165% of that of the female skull. Condylobasal length of the male skull (the same series) M48. 39-5 1.8 mm, that of females is 40.3-M43.39-45.8
the

is

45.5(V.

mm

Popov, 1947).
Extent of individual variation in skulls
variability is of
is

quite considerable in

general, particularly in characters of general size; however, this

"normal" character and does not

attain the ampli-

tude found in

some

especially variable races of weasel. Zygomatic

width

is

strongly variable, width of skull above canines, and

which may be longer and wider or shorter, sharper or weaker. All of this leads to the
particularly in the postorbital constriction
'^Age and sexual variation in the ermine skull was the subject of several special
investigations;
it

certain degree, as a
details, see

was better studied than other species of the family. It serves, to a model for the genus. Very general information is given here. For

Yurgenson, 1933; Stroganov, 1937; V. Popov, 1943, 1947; O. Petrov,

1951; Morozova-Turova, 1961.

1002
fact that skulls within

one race may be more elongated or wider

in description
is

and, in general, obviously differ in their general appearance. This

has repeatedly led to misunderstandings

less than in weasel,

of separate

forms. Geographic variation of the ermine skull

insignificant,
great.

much
Geo-

and these differences are never so

in sexual

graphical changes are not of the degree seen

668

dimorphism.

from the corresponding bone of other species of the genus in that its anterior end is not curved upwards in the form of a hook, and it has a greatly elongated S-shape form. The base of the bone is swollen, with a rough surface, and the anterior end is broadened, with a narrow spoon-shaped depression. A narrow groove extends along the lower surface of the distal half. Age differences in the structure and dimensions of the bone are considerable. In adults (sexually mature), it is larger and heavier. In adult ermines of Volzhsk-Kamsk territory, its length is 23.7-M25.527.7 mm; weight is 0.033-M0.041-0.048 gm; in immature (young and subadult) its length is 22. 2-M23. 5-24.3 mm, weight is 0.016MO.025-0.32 gm (V. Popov, 1947). Dimensions of ermine are variable; however, this variability is "normal", and its extent is not so great as in the weasel. Body length of males (75) is 187-325 mm, of females (47), 170-270 mm; tail length of males is 75-120 mm, of females, 65-106 mm; length of hind
differs

The OS penis

mm, of females, 37.0^7.6 mm; height of mm, of females, 14.0-23.3 mm. Condylobasal length of male skull (213), 39.3-52.2 mm, of females 35.7-45.8 mm; zygomatic width of males 21.0-30.6 mm, of females 18.2-24.2 mm; interorbital width of males 9.1-13.2 mm, of females 7.9-10.9 mm; mastoid width of males 19.1-26.3 mm, of females 16.8-21.7 mm.
foot of males
is

40.0-48.2

ear in males

is

18.0-23.2

Weight of males

to

258 gm, of females

to

180 gm, usually

much

less' (V.H.).

"According

to

acterize ermine of
series; Reichstein,

West Europe. Here, only

Stroganov (1962) data for ermine of our country. They also charindividual animals (judging from a large

1957) with perhaps slightly larger measurements (interorbital width

of males 13.7, females 11.8

the western

mm; zygomatic
e.

width of females 24.6). The given ampli-

tude and dimensions also applies to the small form of ermine from the highlands of

Some measurements given by Novikov mm, one condylobasal length of females 47 mm and others). American ermine may be somewhat larger than European (body length of males 235-340 mm, females 190-290 mm: Hall and Kelson, 1959).
and central Alps (M.
minima).
(1956) are not fully accurate (body length 160-380

1003

Systematic Position

The

relationship of ermine to affiliated species of the genus

in the description of

was

reviewed

weasel (see above) and M. altaica

(see below), with

which

this species is closely related.

The most

fundamental difference
is

lies in the structure

of the os penis which

apparently, important in a biological sense. In contrast to weathe

ermine must be considered as a more "specialized" or weakening of "infantile" features characteristic of weasel, are probably connected primarily
sel,

rather "advanced" form, although the

to larger general

measurements of ermine (allometry). The placing

of ermine at the beginning of the entire series of species of the

genus, as

is

usually done, has no foundation (V.H.).

Geographic Distribution
Arctic and boreal zones of the Old and

New

Worlds.

Geographic Range

in the Soviet

Union

This represents the basic and greatest part of the species range and
occupies the major part of the territory of the

USSR.

The northern border of

Taimyr,
i.e.

the range constitutes the coast of the

Arctic Ocean, to which the ermine reaches even in the northern

to 77

N.

lat.

It is

also encountered on the small is-

lands near the mainland

in the

southern part of the Kara Sea on

the Shokal'sk, Olen'em, Sibiryakova, and

Dixon

islands,

and prob-

670

on Begichev and Salkai islands and islands of the Lena delta. It is known from Bol'shoi Lyakhovsk and Kotel'noe and, apparently, is found in other islands of the Novosibirsk archipelago. It is absent on the remaining islands of the Arctic (occurrence on Baigach is entirely
ably Beloi, Nordenskjold; in the Laptev Sea
possible). In the east, the border of the range passes along the

Bering, Okhotsk and Japanese seacoasts and inhabits Karagin

two of the northern Kuril Islands (Paramushir and Shumshu) and from the South Kunashir. Communications on its occurrence on Iturup (Klumov, 1962) not confirmed by new data (V.G.
Island,

Voronov).
Islands.

It

is

encountered on Sakhalin and

frontier,

all

of the Shantar

The western

from the Barents Sea

to the

Black Sea,

forms the western border.

1005

The southern border of

the range in the

European

part of the

USSR
and
in

and

Kazakhstan extends into the steppe and semidesert, part even the desert zone. In the west, between the state
in

frontier

the Black Sea coast and

and the mouth of the Dnepr, the ermine is met with up to is found even on Tendrov Spit. The erabsent

mine
in the

is

on

the
is

Crimean Peninsula (information of

which
is

Kalenichenko, 1839,

erroneous) and eastwards from the Dnepr

directly adjacent to the

narrow steppe
seas.
It

belt,

Black
at

and Azov
the

exists

immediately around Askaniya Nova and

Don, and perhaps, even below Taganrog and along the northern shore of Taganrog Gulf of the Azov Sea. Farther to the east, the ermine is quite common in the delta of the Volga, is also found farther to the southwest (Mikhailovka, former Biryuche-Kosinsk region, Yasta) and, apparently, is encountered along the Kuma, at least at its mouth. However, there are no prethe
cise data

mouth of

on the southern limit of the ermine's range

at the

in the steppes

of the Cis-Caucasus and

present

moment

the boundary to a

significant extent is conditionally accepted as passing from the mouth of the Don to the mouth of the Kuma. Ermine exists in the Main Caucasus in the El'brus massif, where it is recorded from a series of places from the source of the Baksan to the source of the Cherek (Kabardino-Balkhariya) to a height of from 2,500 to 3,200 m above sea level, in Teberda preserve'^. The inhabited region in Kabarda is, apparently, cut off from the general range of the species. From the mouth of the Volga, the boundary passes eastward along the coast of the Caspian Sea to the mouths of the Ural and Emba and even somewhat farther south. Thence, extending around the northern Chink of the Ustyurt from the north, the southern

boundary of the range extends

Aral Sea, continues along
its

to the

northwestern shore of the

northern shore including Aral'sk

"Data of Korneev (1941), Tembotov (1960) and Basiev (1962). Ermine was discovered credibly in Teberda in 1934 (Korneev, 194) and Kabardino-Balkariya only
in

1959 by V. Dmitriev.

Up

to the

end of 1961, 4 individuals only had been trapped.

this finding did

Earlier, the only

known

data were those of Satunin (1915) about the capture of animal

in the El'brus in the

beginning of the century. Later, when

not receive

confirmation, the occurrence of ermine in the Caucasus was rejected or at least not
all data found in the monograph of Vereshchagin (1959), even those concerning the mouth of the Kuma, were placed in doubt is of some significance for the general problem of the ermine in Caucasus in the last decade since its absence was not accepted either. Up to the present time the ermine has not been found in the Caucasus preserve, although it apparently lives there.

confirmed. The fact that

1006

Karakum and extends

a tongue to the south that includes the lower

Syr'-Dar'ya and descends as far as Kzyl-Orda. Thence, the boundary abruptly ascends to the north, passing through the southern
foothills of the Ulutau, traverses the Sarysu, nearly in its

middle

course and goes on to the Kyzylrai mountains (a little north of the mid-part of [LakeJ Balkhash, 75 E. long.) passing around them

from the south

as well as

Ayaguz

(Sergiopol').

From

there the

border line turns southwards to the western part of the Alakul'sk

depression, leaving
it

at the

range boundary.
in the following

The range

in

Middle Asia may be outlined

form. In the desert lowlands and sands south of [Lake] Balkhash

and along the rivers flow into the lake, the ermine is absent. It expanse between AlakoF and Dzhungarsk Alatau and in this system, between the Dzhungarsk Alatau and Zailiisk Alatau and along the entire Tien Shan system to its western extremity, including the Kirghiz (Aleksandrov) range and, probably, Talas
exists in the

Alatau. Ermine
671

is

absent in the Karatau.

is

The species described

sent.

found
its

in

Fergana and

in the

Pamiro-

Alaisk system. Details about

distribution here are almost ab-

Apparently, however, the ermine occupies the whole Pamir,

Zaalaisk and Darvazsk ranges, and probably Peter the Great range.

Concerning the Gissar, Zeravshan and Turkestan ranges, there are no data. Apparently, ermine does not occur south of Tadzhikistan in the region of low desert mountains. Eastward in Middle Asia,
is everywhere distributed to the state frontier.^" Throughout the distance from the Alakul depression all the way to the mouth of the Ussuri, the range extends southwards to the state frontier and beyond it. Beside, it is possible that ermine is absent in the Amur Valley and adjacent places between the mouths of the Sungar and Khabarovsk. In Ussuri Territory, the ermine is absent in the western part along the Ussuri and the lower courses of its tributaries, and along

the ermine

the coastal zone northwards to a region

somewhat north of

the

^"Range according to data of Kalenichenko, 1839; A.M. Nikol'skii, 1891; 1907; Dinnik, 1914; Satunin, 1915; Orlov and Fenyuk, 1927; Adlerberg, 1935; Ognev, 1935; Rozanov, 1935; Flerov, 1935; Heptner, 1936; Dubrovskii, 1940; Heptner and Formozov, 1941; Vereshchagin, 1947, 1959; Kuznetsov,
Byalynitskii-Biruli,

1948, 1948a, 1952; Shereshevskii and Petryaev, 1949; Heptner, Turova and Tsalkin,

1952; Korneev, 1952; Sludskii, 1953; Bannikov, 1954, Podkovyrkin, 1958; Afanas'ev, 1960; Sokur, 1960; Tembotov, 1960; Baziev, 1961; Stroganov, 1962 and other sources.

1950; Bazhanov, 1951,

1007

mouth of

the Samarga. In this region, the range of the ermine

extends along the main axis of the Sikhote-Alin' southwards approximately to 4430' (the latitude of the southern extremity of
[Lake] Khanka). In the western half of the territory from the
to Sikhote-Alin', the

Amur

the middle Bikin in the

region^'.

boundary of the range apparently passes to form of an arched line from the Khabarovsk

Geographic Range outside the Soviet Union

Occupies Europe from the shore of the Barents Sea southwards to the Pyrenees and Alps inclusive. It is distributed throughout Romania; in Yugoslavia it is only found in the north-southward to
a line passing

from the Danube

at

the junction of Bulgaria and

Yugoslavia

to the Istrian Peninsula

(Riek

formerly Fiume).
To

It is
it

absent in Bulgaria, Albania, Greece and Turkey.

the west

occurs to Ireland and the Hebrides.

In Asia, outside the limits of the

USSR,

the distribution of the

ermine

is

very poorly known.

It

occupies the extreme eastern

montane regions of Afghanistan, Chitral, Kashmir and the region the southernmost place of occurnortheast of Peshawar (Hazara rence in Asia); it apparently lives in the Karakorum and in the western extremity of the Kunlun (the southwestern montane parts of Kashgariya); along the Tien Shan system it occurs eastwards at least to Hami. In the Mongolian Republic, the ermine is distributed everywhere, apparently, except flat semidesert and desert places; it is in the Gobi and Mongolian Altai, Khangai, Kentei and in the foothills of the Khingan. It is distributed in northeastern China (former Manchuria), and according to some data, "everywhere" (Baikov, 1915), but more accurate information is absent. Undoubtedly, it lives in Great Khingan and, probably, in the northern montane regions (Lesser Khingan, Il'Khuri-Alin'), and perhaps also

North Korea. Its distribution in Japan includes at least Hokkaido and Honshu (Hondo). The reference concerning its occurrence in Algeria (whence a
in the east. Possibly
it is

found

in

separate form, M.

e.

algirica,

was described)

is,

to a great degree,

^'In Primor'e Territory, the ermine occupies the highest montane of Terneisk, Krasnoarmeisk, Tetyukhinsk, Kovalerovsk, Chuguevsk and Ol'ginsk regions. These, as well as data given above for Ussuri Territory, were communicated by Bel'skii

(Vladivostok). According to

Maak

(1861), ermine occupies the Ussuri valley.

1008
doubtful, especially
if

we

take into consideration that in Europe,

i.e.

the ermine does not extend south of the Pyrenees and the Alps,
that this

form

is

isolated in

its

distribution.

It is

entirely probable

that this

name

applies to the large southern weasel (M.

.*

673

it is considered by some authors (G. Allen, 1939). North America, the range occupies the entire Arctic archipelago, including Ellsmere Land (Grant), northern Greenland (northernmost point inhabited by the species) and eastern Greenland southwards a little to the south of 70. The southern border of the range in the mainland, north of which ermine is encountered everywhere, forms a quite complicated line, in the west proceeding across middle California, approximately at 40 N. lat. including the northern half of Nevada, almost all of Utah, the greater part of Colorado (except the eastern) and northern New Mexico (the southernmost point inhabited by the species is about 34 N. lat.). Thence, the border sharply rises to southwestern Alberta and southern Saskachewan, and then directs itself southeastwards to North Dakota and northern Iowa, then westwards across the southern ends of Lakes Michigan and Erie, to the Atlantic coast and Chesapeake Bay. It is met with in Newfoundland, on islands off the west coast of Canada, on Kodiak, on Unimak in the Aleutian group (but not on the others) and, possibly, on several other islands in the Bering Sea which are located near the Alaskan coast

numidica), as
In

(no indications for the Pribilovs) (V.H.).

Geographic Variation
In vast area of
its

range, the ermine reveals

is

some geographic

vari-

ation which, however,

not great.

Its

amplitude, in a morphologiin the weasel,

cal sense does not attain the

is

dimension observed
It

and

not more than that in other comparatively slightly variable spe-

cies of the family

and genus.

mainly bears a clinal character.

The winter coat

and
in

(its

density, length), the color of the

summer

fur,

part the general dimensions, are chiefly those that vary

geographically.

Geographic variation of the ermine is still insufficiently studand representative of this, it is, apparently, exaggerated. Within the last decade, in reviews of our fauna, 10 subspecies were at first
ied,

*In Russian original, erroneously as e

Sci. Ed.

1010
accepted (Ognev, 1935);
later,

although with reservations

13
less.

(Novikov, 1956). In recent times, a tendency to reduce the number of forms has been noted (Morozova-Turova, 1961). Apparently,
the actual

number of

realistically characterized races

is,

still

Several foreign investigators accept

many forms

21

for the for

World (Ellermann and Morrison-Scott, 1951) and 20

(Hall, 1945; Hall
is

Old America

and Kelson, 1959), where the extent of the range

for our

much

smaller than in Eurasia.

The following forms may be provisionally accepted

Stroganov, 1962, with modification).

country (characteristics mainly after Morozova-Turova, 1961 and

1. Northern ermine, M. (M.) e. erminea Linnaeus, 1758. Dimensions moderate or small. Facial part of skull is relatively short and broad. Condylobasal length of male skull 43-49 mm. On Kola Peninsula.

Outside the
the

USSR Scandinavian

Peninsula.

Kola ermine are poorly known and the actual relationship of

name

to this race requires confirmation.

2. Middle Russian ermine, M. (M.) e. aestiva Kerr, 1792. Dimensions moderate. Color of summer fur dark-tawny or chestnut. Individual variation in color is insignificant from light-reddish to dark-tawny

tones.

Body
is

length of males (57),

215-M248.7-278 mm, of females

195-283 mm; tail length of males,78-M92.4-105 mm, of (23) females, 60-86 mm; length of hind feet of males 41.5-M46.2-55

mm.

mm,

Condylobasal length of skull of males (76), 41.0-M46.8-52 of females (20), 39.8-M42.0-45.2 mm; zygomatic width of males, 21.5-M26.0-27.8 mm, of females, 21.2-M22.5-25.3 mm; interorbital width of males, 10. 0-M12. 0-13.0 mm, of females,

M10.2 mm*; mastoid width of male

of females,

18.0-M21.1-24.5 mm 54.2-58.1% of condylobasal length. European part of the USSR, except Kola Peninsula. Outside the USSR, in Middle and West Europe. 3. Caucasian ermine, M. (M.) e. teberdina Kornejev, 1941
balcarica).

19.0-M22.9-26.2 mm, Zygomatic width constitutes

skull,

(syn.

Dimensions small. Color coffee- or reddish-tawny.

*Only mean value
in

Russian original

Sci.

Ed.

1011
674

mm; tail mm; length of hind females, 27-30 mm. foot of males Condylobasal length of male skull, 43.0 mm, of females, 37.8-39.3 mm; zygomatic width of males, 22.6 mm, of females, 20.020.6 mm; interorbital width of males 9.8 mm; of females, 18.4-18.6 mm. (Baziev, 1962).
Body
length of males 217

mm,

of females, 175-190

length of males, 84

mm, 40 mm, of

of females, 57-61

Northern slope of middle part of Main Caucasus range (El'brus

massif).

Outside the

USSR absent.
known form,
e.

very

little

described as similar in dimensions

(!).

and color to

M. (M.)

ferghanae

The independence of

this

form and its assigned characters require confirmation. 4. Tobolsk ermine, M. (M.) e. tobolica Ognev, 1922 (syn.
ognevi, birulai, martinoi).

Dimensions
aestiva.

large,

on average somewhat larger than

silky.

in the

form

Winter fur
tutes

tall,

dense and

Summer

fur

is

also

somewhat
its

longer, denser and

softer. Skull is large.

Zygomatic width constiskull.

54.6-M57.7-58.1% of

the condylobasal length of

Body length of males (43), 200-M260-300 mm, of females, (35) is 200-M230-270 mm, tail length^^ of males is 50-M90-120 mm, of females 50-M70-100 mm; length of hind foot of males 40-M45-48 mm, of females, 34-M37-42 mm; height of ear of males 18-M20-22 mm; of females, 14-M16-18 mm. Condylobasal length of male skull (45), 43. 4-M48. 3-50.1 mm,
of females, 39.6-M43.0-45.6 mm; zygomatic width of males, 24.8M27.7-30.0 mm, of females, 21.4-M24.2-26.8 mm; interorbital width of males, 10.4**-M12.0-13.2 mm, of females 9.2-M10.812.0

mm,
mm,

postorbital width of males, 9. 6-Ml 1.2-12.1

females, 8.2-M9.2-11.1

25.2

of females,

mm, of mm; mastoid width of males, 21.2-M23.61 9. 3-M2 1.0-22.2 mm.

and
Altai,

In western Siberia, eastwards to the Yenisei

and

in

Kazakhstan.

Absent outside the USSR.

^^The

minimum

of this measurement (Stroganov, 1962) strongly differs from that

(p.

given previously according to the same author

mistake
is

668) and, apparently, here the

used, which cannot be found. form of the verb to bungle Sci.
is

self-inflicted*.

*In the Russian original, the

It

word "Kroetsya"

may be

a misspelling of "kropatsya", the reflexive

Ed.

**In Russian original, erroneously as 104

Sci. Ed.

1012

For

this

form, in contrast to the form aestiva the following are

considered characteristic: particularly large dimensions, widely

diverging zygomatic arches and several other craniological fea-

(Ognev, 1935; Morozova-Turova, 1961). According to Stroganov (1962), only characteristics of fur distinguish the Tobalsk ermine from the form aestiva.
tures
5. Altai ermine, M. (M.) e. lymani Hollister, 1912. Dimensions moderate. Fur less dense than in M. e. tobolica. Color in summer fur with weakly developed reddish-brown tones, sometimes this tinge disappears completely. Skull similar to that of M. e. aestiva. Distance between zygomatic arches relatively narrow (zygomatic width constitutes on average about 55% of condylobasal length). Body length of males (10), 218-M258-295 mm, of females,

185-M220-260 mm; tail length of males, 45-M75-105 mm, of 40-M60-90 mm; length of hind foot of males, 37-M4448 mm, of females, 32-M36-42 mm.
females,

of females, 41.2-M43.7-45.2

Condylobasal length of male skull (14), 44.0-M48.4-49.4 mm, mm; zygomatic width of males, 24.4-

M27.3-28.7 mm, of females, 20.8-M24.4-26.2 mm; interorbital width of males, 11.8-M12.2-12.6 mm, of females, 9.9-M11.312.0

mm;

postorbital width of males, 10. 0-Ml 1.4-12.8

mm,

of

females; 9.0-M10.2-1 1.7

675

mastoid width of males, 21.3M23.6-25.3 mm, of females, 19.2-M21. 1-23.0 mm. Weight of males (5), 145-M191-247 gm, of females (4), 106149 gm (Altai preserve, winter; material from ZMMU).

mm;

Mountains of southern Siberia eastwards

Sayan).

to

Baikal (Altai,

Outside the

USSR in

contiguous parts of the Mongolian

is

Republic. In the Gobi Altai and southern parts of Khangai and the

Mongolian
6.

Altai, apparently,

another form.

e. kaneii Baird, 1857 (syn. naumovi, sibirica, kamtschatica, kanei, digna; sometimes the name arctica was, and still is applied, to this form). Dimensions moderate, less than in M. e. tobolica. Color of summer fur relatively light, with brownish-yellow

East Siberian ermine, M. (M.)

orientalis,

tinges of various intensities.

Zygomatic width constitutes about

56% on
(7),

average of condylobasal length of skull.

Body length of males (22), 213-M260-325 mm, of females 176-M212-222 mm; tail length of males, 70-M86-100 mm,

1013

67-M72-77 mm; length of hind foot of males, 40of females, 33-M38-43 mm; height of ear in males, 20-M21-22 mm, of females, 18-M 19-20 mm.
of females,

M4348 mm;

of females (23), 3 8. 9-M40. 9-44.3

Condylobasal length of male skull (24), 44.0-M46.0-49.2 mm, mm; zygomatic width of males,

24.0-M26.2-28.8 mm, of females, 20.0-M22.3-25.2

orbital

mm;

inter-

width of males, 10. 1-Ml 1.4-13.2

postorbital width of males,

mm,

of females, 8.8-

M9. 5-10.0 mm;

10.0-Ml 1.0-12.5 mm,

of females, 8.7-M9.7-10.9

mm;

mastoid width of males, 20.1-

M22.5-24.7 mm, of females, 18.0-M19.2-20.2 mm. Weight of male (8), 102-Ml 66-253 gm (Kamchatka, summer; material from ZMMU). In eastern Siberia and the Far East including Kamchatka, except the

Amur

area and Ussuri territories, Trans-Baikaliya and

Sayan.

Absent outside the USSR.

This form
is

not sharply distinguished, differing from western

Siberian tobolica only in slightly lighter color and

dimensions. Differences can only be noted

close to M.
7.
e.

in

somewhat smaller series. The form is

identical with
it.

arctica from Alaska and

e.

may be

Karagin ermine, M. (M.)

karaginensis Jurgenson, 1936.

Dimensions small, significantly less than preceding forms. Color of summer fur light-chestnut. Body length of males, 220-230 mm; tail length 75-80 mm; length of hind foot 37-39 mm. Condylobasal length of male skull, 40.5-M42.7-43.7 mm; zygomatic width, 21.6-M23.0-23.7 mm; interorbital width, 9.8MlO.1-11.0 mm; mastoid width, 19.3-M20.3-21.6 mm. On Karagin Island along the eastern coast of Kamchatka. Absent outside the USSR. This form is poorly known. Its independence must be confirmed in new material. It is possibly related to the form kaneii. 8. Trans-Baikal ermine, M. (M.) e. transbaikalica Ognev, 1928
(syn. baturini).

Dimensions

relatively small.

Summer
stitutes

fur short

with relatively narrow interzygomatic area

and sparse, dark-brown in color. Skull small zygomatic width con-

on average 51.6% of condylobasal length of skull. Body length of males (14), 225-242 mm, tail length 80-M85.7length of hind foot,

90

mm,

31-M40-45 mm.

1014

Condylobasal length of male skull (94), 39. 3-M43. 6-46.1

mm,

of females (26), 35.7-M38.2-41.7 mm; zygomatic width of males, 2 1.0-M23. 2-24.5 mm, of females, 18.2-M19.5-23.7 mm; interorbital width of males,

9.1-M10.8-11.6

mm,

of females, 7.9-M8.7-

10.0
676

mm.

Postorbital width of males, 9. 3-Ml 1.0-12.0

mm,

of females,

8.3-M10.1-10.7

mm;

mastoid width of males, 19.1-M21.0-22.1

of females, 16.8-M18.1-21.8 mm. Weight of male (9), 101-M134-174 gm (Barguzin preserve, winter; material from Z[oological] M[useum of] M[oscow]

mm,

U[niversity]).
In Trans-Baikaliya,
lands.

Amur

area, Ussuri Territory, Shantar Is-

Outside the
lian

USSR

probably

in the eastern part

of the

Mongo-

Republic and northeastern China (former Manchuria).

This form
is

close to the Altai,

M. (M.)

e.

lymani and possibly

all

both

may comprise one

has priority.

form, which occupies

of southern

montane Siberia and above-mentioned

parts of the Far East.

The

name lymani
9.

Fergana ermine, M. (M.)

e.

ferghanae Thomas, 1895 (syn.

schnitnikovi)

Dimensions small. Color of summer fur very

rarely with a film of red.

light,
is

straw-brownish or grayish,
short and soft.

The coat

On

the neck

occur light spots, sometimes forming a collar. This

winter or

is

the lightest

colored form of our fauna. Individual animals do not turn white in

become only

partially lighter.

M75-82 mm;

205-M259-300 mm; tail length 7040-M41-42 mm. Condylobasal length of male skulls (9), 40-M43-45 mm;
Body
length of males, length of hind foot,

zygomatic width, 22.0-M23.3-25.5 mm; interorbital width, 9.510.4-1 1.0 mm; mastoid width, 19.0-M20.6-22.0 mm.

Weight of males in summer (12), 121-M145-211 gm, of females (10), 60-M72.5-86 gm (material from Z[oological] M[useum of] M[oscow] U[niversity), from Tien Shan, mainly
Zaillisk Alatau).
In

montane Tien Shan and Pamir-Alaisk system.

Outside the

USSR,

parts of the range in Afghanistan

and the westernmost parts of Tibet; found Shan China.

in adjacent parts

and India of Tien

1015
It is a well differentiated form, to which the ermine of the Gobi Altai and adjacent parts of the Mongolian Republic imongolica, Ognev) apparently belong.

The number of subspecies of ermine described from

the range lying outside the boundaries of the

parts of

USSR,

is

great and

evidently exaggerated, particularly for America.

graphic 1904 England;

Hebrides; 5)

The following forms are usually mentioned: \) M. (M.) e. hibernica. Thomas et Barrett-Hamilton, 1895 Ireland; 2) M. (M.) e. algirica Thomas, 1895 Algeria (see reference to this form in the section "Geo-

distribution"); 3)
4)

M. (M.) e. stabilis Barrett-Hamilton, M. (M.) e. ricinae Miller, 1907 islands of the M. (M.) e. minima Cavazza, 1912 Switzerland; 6)

Japan; 7) M. (M.) e. mongolica e. nippon Cabrera, 1913 Ognev, 1928 Gobi Altai (apparently, a synonym of ferghanae); Alaska, northwestern parts 8) M. (M.) e. arctica Merriam, 1896 of Canada, Arctic archipelago, except Baffin Land; 9) M. (M.) e. polaris Barrett-Hamilton, 1904 Greenland; 10) M. (M.) e. semplei Sutton et Hamilton, 1932 Baffin Land and adjacent part of mainland; 11) M. (/.) e. richardsonii Bonaparte, 1838 Newfoundland, Labrador and nearly all of Canada except the territories occupied by the forms named above; 12) M. (M.) e. cicognanii Bonaparte, 1838 region north and east of the Great Lakes; 13) M. (M.) e. bang si Hall, 1944 region west of the Great Lakes; 14) M. (M.) e. murica Bangs, 1899 southwestern extremity of the range

M. (M.)

of the species in America (Nevada, Utah, Colorado and other states);

15)
677

M. (M.)
e,

(^.)

e. kadiacensis Merriam, 1896 Kodiak Island; 16) M. anguinae Hall, 1932 Vancouver Island. From a small part

of the range the southern part of British Columbia, the state of Washington and western Oregon described forms are; 17). M. (M.) e. fallenda Hall, 1945; 18) M. (M.) e. invicta Hall, 1945; 19) M. (M.) e. gulosa Hall, 1945; 20) M. (M.) e. olympica Hall, 1945; 21) M. (M.) e. streatori Merriam, 1896; all have a limited or extremely limited distribution. Described from various islands of the Alexander archipelago and in part from the adjacent coastal mainland are; 22) M. (M.) e. alascensis Merriam, 1896; 23) M. (M.) e. initis Hall, 1944; 24) M. (M.) e. salva Hall, 1944; 25) M. (M.) e. celenda Hall, 1944; 26) M. (M.) e. seclusa Hall, 1944; 27) M. (M.) e. haidarum
Preble, 1898 (V.H.).

1016

Biology
Population. Ermine belongs

among
its

the

nivores. Within the boundaries of

sity are,

range,

number of abundant carits numbers and den-

however, distributed very unevenly.

Indices of prepared skins, even those calculated per unit area

(usually per 10 km^) by no
the species population.

means reflect the actual condition of They depend, to a very strong degree, on

the extremely different intensiveness of harvest, the degree of

control on animal stocks, and finally, on the condition of the population itself in the given trapping season.
It is

established that the

catch and skin preparation increase with deficiency of ermine foods,

but

is

not proportional to

its

numbers

(S.

Severtsov, 1941; Teplov,

1952). This takes place due to the greater ease with which hungry

animals are caught

in baited traps in a

year following one with

abundant reproduction.

The ermine

is

the

most numerous

in the forest-steppe regions

of western Siberia and northern Kazakhstan. In second place fol-

low the forest-steppe regions of Bashkir ASSR, Tatar

ASSR

and

of the middle Volga region, and also the southern regions of

Krasnoyarsk Territory, taiga and tundra regions of western Siberia and Komi ASSR. To the east, west, and south, the ermine is met
with significantly more rarely.
southern Ukrainian
the
It is

more frequent
it

in the steppes of

SSR

and

in the southeastern
is

European

part of

USSR

(P.B. Yurgenson). Here,

restricted almost exclu-

sively to the deltas and floodlands of the great rivers.

For nine years (1924/25-1933), the average yield of skins per

10

km^was

as follows: Bashkir

ASSR 1.82,
is

Siberia 1.07, Tatar

ASSR 0.78,

Ural 0.85,

Severnyi Territory0.60. In other parts

considerably lower.

of the range, abundance of ermine

Of course,

these figures are connected not only with the abundance of ermine,

but also with the development of the harvest in these years. At the
present time,
it

has fallen significantly everywhere. At the same

time, in a series of the

dant,
its

main regions where the ermine was abunnumbers fell sharply, especially in the forest-steppe of western Siberia and northern Kazakhstan, due to the deterioration of food resources, destruction of shrubby growths along lakes, worsening of the hydrological regime, etc. Habitat. Habitats of the ermine are sufficiently variable.
Nevertheless, in different geographic zones,
it is

closely associated

with near-water biotopes.

1017
In the tundra zone, the

ermine prefers the banks

It

'of rivers,

riparian

meadows and

thickets of bushes.

also lives along the

slopes of valleys and in rocky places.

typical for

The

latter are particularly

montane tundra.

It

also willingly holds to the fringe of

it prefers most of all the whole of the banks and floodlands of rivers and creeks, the shores of forest lakes, floodlands and marshy sedge meadows with shrubby

the forest-tundra. In the forest zone,

thickets and

hummocks,

etc.

Forest massifs are untypical habitat

it

for the ermine, although here

is

met with almost everywhere, but

settle-

not often. In forests, old cluttered burned areas and glades, the
678

edges of forests and shrubby areas (especially those near

ments and arable lands), but deep within forest massifs it prefers cultivated fir and alder as well as the forest meadows and old haystacks. It does not avoid settlements and gardens near them. In winter, it often searches for food in grain stores and heaps of straw. Reduction in area of cut-over areas and burns resulting in age
changes
in the

cover of forest vegetation covering them, leads to

a sharp decrease in the

number of ermine
it

in the forest

and

to

decrease in the area inhabited by

(for instance,

100 times). As

ermine occupation which are under surviving stands, whence teeming with life. These are original favorable conditions ermine spread out into other stands. Flooded
a result of this process, the greatest density of
in fir forests is

found

in the cultivated fir plantations

fir-groves

when they cover

the

whole

forest massifs, favor

anew

the settlement of animals in newly-appearing clearings and burns

from many points (Yurgenson, 1959). In the forest-steppes and steppes, ermine clearly avoid open expanses of steppe and restrict themselves to floodland meadows
with osier thickets along the valleys of rivers, along steppe clearings and their edges, in birch-groves along

swamps,
It

in

hummocky
It

swamps and

in

weedy

thickets around steppe lakes.

is

also

encountered along steppe ravines and gorges.

cities.

often appears in

dwellings not only of small settlements, but also the outskirts of

ermine occupies rock slides with pika colonies, and dry alpine meadows inhabited by the narrow-skulled
In the Altai mountains,

vole [Microtus gregalis]. Here,

plastic than sable
in

it

more willingly
It is

lives in places
is more met with

with shallow snow cover. In selecting stands, the ermine

and Siberian weasel.

in

more

rarely

dark coniferous taiga than

thin,

sunlit stands.

On

coastal

1018

and catches water voles by unfrozen springs; in montane taiga, it is an inhabitant of rocky sections and open elans*. In winter on montane balds and bushy tundra; it sticks
cliffs
it

lives in rock slides,

to stands with

slides inhabited

willow ptarmigan and lives for a long time in rock by pikas (Dul'keit, 1956). In the mountains of
its

southern and southeastern Kazakhstan places of

include shrubby thickets, apple, juniper and
fir

occupation

forests

and highpre-

land fescue steppes (Sludskii, 1953).

fers floodlands to other types
its

The ermine everywhere

it

and without them,

spends

much

of

time near water. In Tatariya, in flooded areas, the density of

is

ermine

six times higher than in

plakorn* dry regions (Aspisov

and Popov, 1940). Food. According

rodents (in the

to feeding type, the

ermine belongs among

its

animals with a limited assortment of food. In

food mouse-like

broad sense) predominate, but

food

in contrast to the

weasel which almost exclusively feeds on small voles, a considerable place in

its
is

occupied by the larger rodent species

water vole,

common
in their

hamster, pikas and others, which the ermine

overpowers

burrows and which the weasel cannot master

or with great difficulty.

More
birds,

or less constant, but of secondary significance, are small

fish

and shrews. More rarely there are amphibians, reptiles (lizards), and insects. All of these secondary and rare foods in the diet of the ermine, and also berries in Lapland (Nasimovich, 1949), are a sign of food deficiency, deprivation, and simply starvation. In the Ob' preserve, 156 cases of bird nests and sometimes

destroyed by ermine were recorded during four years.

Of

these in

60 cases
16

it

was

nests of the pied flycatcher, in 59

great

tit,

in 8

spotted

flycatcher, in 6

wryneck, and

starlings, in
in

white wagtail (Karpovich and

Sapetina, 1958). In years with high

and prolonged flooding, the number of cases grows since bird nests are destroyed on the ridges and islands during inundation. Here, nest destruction comprises 14%, while outside the flood zone 0.5%.
In the

ermine of floodlands and banks of watercourses, besides

field

the water vole and gray vole [Clethrionomys rufocanus], the root,

common and

voles [Microtus oeconomus, M. arvalis, M.

680 agrestis] predominate; in forest areas, various species of red-backed voles; in tundras,

lemmings
Sci. Ed.

also.

Ermine

rarely attack the

commercial

*Loca{ word?

'

1019

Vi

3 ^uS U ON ^
=

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"
7^

11^1

CQ .
'

^
S

^
j2 :S

^
-

I
'^

S 2

^ > ^^ -

23

2 5

15^111111^
vo
j:^

III
-

g
'-

N
>

2 --

- I

00

"^ u->

tn

I
I

d
I

^O'^t^
rn

d d

lo

^"Q .^^
^J

g-c;

'^ 0\

Tt "*

.S o^

^i
^

2 II

III

S
V S
"tt

>
i2

> >
l-l

^
I

2 II

:2^

^
00

" J > W -

y-

5
I

II

II ^

III

oo

m
00
I
I

00

U
.5

f*^ <?^

OS

I
I

-I

fN

<)

-1

00

"
S > 6

JC

.^>
1 1

00-- _
QJ

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00

'3

U D t

CQ <;


1020

However, cases of ermine feeding on muskxats, probably young, have been recorded. Geographic variation in nutrition of ermine is not great. Variations in the predominant food composition in different years are significant. They depend on the numerical composition (yield) of the different species of mouse-like rodents. These variations are partially presented in Table 63 (data from Komi ASSR, Pechora and Kirov district), and for Pechora-Ilych preserve over 12 years in Table 64. In Laplandia, in years with abundant voles and lemmings, an ermine caught four animals in one hunt (Nasimovich, 1948). It
animals and birds
rarely with success.

even

more

eats

no more than two and

it

in a

famine year, one

satisfies

it.

With
it

an abundance of prey,

consumes only

the brain or leaves

more animals than it is able to eat. The daily food norm is about 50 gm, i.e. about 25% of the live weight of the animal. The ermine usually makes a stockpile of food. Found among its "stores" were up to 5 common
untouched on the
trail,

killing significantly

hamsters, 5-8 water voles, up to 10 small mouse-like rodents and

4 shrews

at

one time,

etc.

(Zverev, 1931 and others).

Home 12-15 cm

range. With the formation of

snow cover deeper than

or more, each ermine occupies a defined

home

range.

680 Table 64. Nutrition of ermine in the winter period in Pechora-IIych preserve over
a 12-year period

(%

of occurrence)

Type of food

Riinge of fluctuation
in

occurrence

Average for 1938/39 to 1950/51 47.9 24.2 7.8

11.8 1.9 0.7
1.5

Voles

072.7
5.966.9

Shrews Tetraonid birds Other birds

Squirrel

013.3 035.8
011.2

White hare
Carrion
Insects

07.4 06.7 07.1

of empty

0.5

Percentage

stomachs

2958
0.5 3.0g

46.8
1.35g

Average weight of stomach

contents
Note.

From

data of the "Nature Chronicle" of Pechora-Ilych preserve.

1021

This area
with repeated

is

divided into a series of ranges of daily activity,

visits

each 3-10 days (Zharkov, 1941). Such ranges

of mingled living animals often overlap each other.

In sparsely treed, northern

of the

home range

varies within limits of

montane Murmansk district, the area 50-90-100 hectares

(Nasimovich, 1948). In the forested regions of the middle belt, its dimensions shift from 10-15 to 100-200 hectares, depending on
food. Here, the range of daily activity at the start of winter equals
681

5-6 hectares each, but by the end

daily activity range varies

it

reaches 100-200 hectares

(Shibanov, 1935). In the floodlands of Tatariya, the area of the from 7.7-8.8 up to 31.5 hectares. The
area of the daily activity range of males
is

considerably greater

than in females: in males 15-20 hectares, average 29.5;* in fe-

males up

to 8 hectares,

average 7.1 hectares. Sometimes, a group

possible that

of ranges are observed, inhabited by 5-12 individuals and alternating with ranges of lower density of occupation.
It is

these are mingled ranges of the dispersing

litter

(Zharkov, 1941).
a

Burrows and
pendently.
It

shelters.

The ermine does not dig

burrow inde-

often uses the burrows and nest chambers of the

rodents
others

chipmunks, and and even more often as a temporary shelter. The nest chambers in the brood burrows are lined with the skins and underfur of mouse-like rodents, rarely only with dry grass. Sometimes, the brood shelters are located in very unexpected and seemingly unsuitable places, for example among logs
it

kills

water

voles, hamsters, Siberian

for the litter young,

piled against the wall of a house in a large village (Kazakhstan;

V.G. Heptner) and without any kind of lining at all. The ermine also lives in old and rotting stumps, under tree roots, in heaps of brushwood, haystacks in floodland meadows, in heaps of straw, in old haystacks, in bog hummocks, in the cracks of vacant mud buildings, in rock piles, rock clefts, and even in

magpie

nests.

It

also occupies the hollows of trees;

it

particularly

often uses these during periods of flooding. Males and females

live separately, but near

each other (Velizhanin, 1931; Zverev,

1931; Yurgenson, 1932; Nasimovich, 1948; Sludskii, 1953). The ermine has no permanent shelters in winter, and uses the
opportunistic ones near
its

hunting places.

It

rarely returns to

its

old place of day

*There
determined
is

rest.

Such

shelters occur under rocks, in the roots

an obvious misprint here, but what the correct values are could not be
Ed.

Sci.

1022
of trees, under logs covered with snow, etc. However, in Yakutiya,
apparently,
it

uses one burrow throughout the entire winter (Belyk,

1958).

Daily activity and behavior. The ermine

to pass that

is

mostly active
it

in the

twilight and night hours of the diel period, but sometimes

it is

comes
in

encountered during the day. The light regime

high latitudes substantially influences the diel cycle of the ermine.

In the period of the polar day, about
this

70%
is

of the occurrences of
often

animal take place

in the night

hours (1800-2400). In autumn

(September-October), when the night

is

becoming darker,

it

encountered during the day. In the

first,

darkest half, of winter

(15 Nov.-l Feb), the ermine most often hunt in the morning and

during the day (morning

night

5; Nasimovich, 1948). During periods of severe frosts activity of the ermine notably
its

11

encounters, day

7,

evening and

decreases. With

available stores of food

it

can remain,

in this

event, for 1-2 days, as well as after a snowfall. During snowstorms

it

is

active (Velizhanin,

1951; Zverev,
it

Nasimovich, 1948). In winter,

day.

1931; Zharkov, 1941; goes out for prey 1-2 times per

In winter, during one excursion, the ermine travels 0.5-8.0

km

(about 3

km

on average). In short

lateral

loops

its

wandering "shut-

tle" route constitutes 1/3 to 2/3 of its daily

journey (Nasimovich, fundamentally a

not able to

1948).

The ermine swims and climbs

terrestrial animal. It

well, but

it

is

can

move

freely in the underground passages

it

of rodents

water

voles, hamsters and pikas, but

is

penetrate the burrows of small voles.

brave and cunning. The prey

is

killed

The ermine is very bold, in the same manner as the

weasel, by biting the skull in the occipital region. In a situation,

where

it

is

cornered,

it

dares to attack humans. In the tents of

it

peoples of the far north

often

makes

itself

an intrusive lodger,

plundering stock of meat, fish and other products.

The movement of the ermine is quick and dexterous, but somewhat fidgety. It is extremely curious, a property which is closely
connected with careful examination of the hunting range when
acquiring food.
682

Seasonal migrations and transgressions. Sedentariness within

the range boundaries
it

is

typical for ermine in winter in years

is

sufficiently

supplied

with

food;

when when unbalanced by


1023
starvation or insufficient food, then
it is obliged to roam widely. Lapland preserve, an ermine roamed an area of 25 km^

In

(Nasimovich, 1948).

Sometimes, the abundance of water voles or mouse-like

ro-

dents and characteristics of their distribution within an area lead to

massive displacement of ermines for considerable distances. Such migrations were noted in western Siberia in 1928 and were compared with mass reproduction of water voles in the northern regions (Zverev, 1931). The mechanism of such migrations remains

almost unexplained.
In a series of localities, moreover, local migrations are ob-

served of seasonal character. Thus, in Voronezh forest-steppes,

ermine

in

winter

move down from highland oak groves

to floodland

black alder forests

levadas* (Severtsov, 1850)** and in spring-

back. In forests of the middle belt,

some ermine move from

the

edges of the forests and shrubby thickets to populated areas

stores, straw piles

grain

and haystacks, feeding on voles and mice con-

centrated there. In the floodlands of large rivers, during the time

of spring flooding, ermine are concentrated in the inundated ridges

where an abundance of food in the form of mouse-like rodents migrating thence from the floodlands are found (Aspisov and Popov, 1940; Zharkov, 1941 and others). In Oka preserve, during time of the spring flood, on unsubmerged islands, five pairs of ermines were found in an area of 36 hectares, which constitutes a density of 278.0 per 10 km^ (Karpovich and Sapetina, 1958). In the Lapland
preserve, in connection with dispersal of forest voles, local migra-

from the pine woods and burns to the fir groves for a distance of 10 to 15 km were observed (Nasimovich, 1948). Reproduction. Reproduction of the ermine is still entirely insufficiently studied. It has one sexual cycle per year. Spermatogenesis in males lasts for five months of the year, and sexual activity for four months (middle of February-first half of June for northern
tions

Kazakhstan). In the nonfertilized females, ovulation

the

is

periodically

repeated monthly. They are capable of being fertilized throughout

whole period of male sexual activity. In adult females, most part takes place just after parturition. Young females are fertilized in summer. During the course of
fertilization for the

pregnancy, there
*

is

prolonged latent period, probably with

Local name? Sci. Ed. ** Misspelled Severtsev in Russian original? Not in

Lit. Cit.

Sci. Ed.

1024
variable duration. After
lasts not

it,

the active

development of the embryos

varies

more than one month. The number of young in a litter

from 2-18, more often

is

4-8; for northern Kazakhstan, the average number of young

estrus usually occurs in

8.7

(Tikhvinskii, 1937; Grigor'ev, 1933; Lavrov, 1944). In Yakutiya,

April-May, but sometimes even in the first embryos are observed easily by eye. Their number ranges from 7-8 up to 12. In Yakutiya, the number of males in a
half of April,
litter is

greater than that of females (Belyk, 1958). In populations

of West Siberia, males constitute

53% and

females

47% (Zverev,
of the popula-

1931). In northern Kazakhstan, males are

tion;

61-65%

on average, there is one female for 1.75 males (Lavrov, 1944). The sex ratio changes by season, reflecting the degree of activity of the different sexes; sex ratio also changes substantially by year, depending on environment conditions of the external environment.
In unfavorable years with insufficient food supply, the

number of

females increases (Lavrov, 1944),

In
in late

ermine of Yamal, two peaks of reproduction are observed:

winter and in summer. In July-August, the main mass of the

current year's females participate in the rut, having attained sexual

maturity at the age of 3-4 months. Adult males also take part in
it

(these born in the preceding year), because the males of the

current year attain sexual maturity only in the following year.

683

After pregnancy which lasts (with the latent period) 9-10

months, the young are born

females

in

April-May. At

that time, those

who have not, for various reasons, participated in the summer rut, come into heat: in them pregnancy is only six weeks, the
young also appearing in May-June (Kopein, 1961). The timing of sexual activity and reproduction
geographic variation, but
April, blind,
this question
is

subject to

has not been studied.

in

Growth, development and molt. The young are born

March-

naked and helpless. Both parents participate in their care (Bekshtrem, 1931; Sludskii, 1953 and others). When rivers overflow onto floodlands, they transfer their young to a safe place. At the age of one month, the still-blind young are already covered with relatively dense fur and have well-developed canines and molars. At the age of 2-3 months, the young attain the size of the
mother, differing only in fur quality (Lavrov, 1944). At the end
of June-July the young are independently capturing food, but the brood still has not dispersed (Zverev, 1931). Young males attain

1025
adult dimensions only in the second year of life. Young females become sexually mature very early at an age of 2-3 months and, probably, are quickly made fecund. Spermatogenesis is not yet observed in young males in their first year of life (Lavrov, 1944).

Two

molts occur in ermine: spring and autumn.

Enemies, diseases, parasites, mortality, and competitors. Red

fox, golden eagle (Nasimovich,

1948) and sable (Baturin, 1923;

Dul'keit, 1929; P.B. Yurgenson) are

among

the

immediate enemies

of the ermine. References to other animals are based on general

considerations, not confirmed facts.
In the

Sayan mountains, the ermine

sable,

as well as the Siberian

weasel occupy successional habitats which are sometimes isolated

from the habitats of

and the

latter

animal occupies the widely

better with sable

distributed primary habitats.

The ermine coexists

than does the Siberian weasel. Dispersal of ermine takes place at

end of November-beginning of December when the sable has its habitats. Concentrations of ermine tracks are observed in places sable and also squirrel have left: floodlands of rivers, burns, and places of heavy snow accumulation (Dul'keit,
the

already determined

1959).

Diseases of the ermine are insufficiently studied. The ermine weakly susceptible to tularemia (Lavrov, 1944); disease of a tubercular character and a type of infectious encephalitis type have been noted (Yurgenson, 1931; Lavrov, 1944). Infestation by tapeworms is not great 38.5% (Lavrov, 1944). The helminth fauna of the ermine is not rich (10 species), and the degree of infestation is usually insignificant. Only skryabingulosis is of substantial pathois

logical significance. In different years, the degree of infestation

reaches
in

50%

of the population, and the number of individual

one ermine

up

worms
is

to 45.

Beside mortality, skryabingulosis causes

a reduction

in

fertility.

Tapeworm

invasion of the ermine

particularly dangerous in poor food years (V.

Popov, 1943, 1947;

Lavrov, 1944).

The extent of longevity in the ermine is unknown. Age determination beyond 2 years by the tooth wear method (Stroganov, 1937) is not confirmed by accurate data and therefore
it

is

not insufficiently founded. In populations of the species, ani-

mals up to one year of age constitute, according to commercial

capture data, from 38.5 to

69%

(average 54.2%), those 2 years

old from

20.8 to 33.8% (average 24.5%) and those above 2

1026
years

from

9.7 to

figures, selectivity of the harvest

37.4% (average 21.3%). In evaluating these must be taken into consideration.

In Tatariya, the quantity of juvenile animals varied over 11 years

from 19.3
1947). In

to 63.9%, constituting an average of 50.8% (V. Popov, Yamal, among the ermine inhabiting the tundra in sum-

mer, juveniles constituted 72.5% of the population as a whole, but

adult, the reproducing part of the population, constituted

27.5%. In

684

young ermine constituted 94.4% of the population, and 5.6% (2,373 specimens). These data speak to the higher adults mortality among adults. Usually, mortality of young is higher. For ermine in the far north it has been shown that average length of life is 1 year, and the period of complete turnover of the population 3 years. Death of adult ermine occurs in autumn, and juvenile females play the main role in increase in the number of ermine in the following year (Kopein, 1961). The argument for increased autumn mortality of adult ermine is not ecologically based; the main since in autumn, the number of mouse-like rodents
winter,

food of ermine
In
is

is

highest.

Murmansk

district, the principal

reason for ermine mortality

starvation (Nasimovich, 1949). In Kazakhstan, ermines die in

great

numbers during the time of great steppe

from snowmelt water
in spring;

fires; in

connection
as a result

with drying up of water bodies in the steppe; during flooding of

the steppe

from hunger

of frozen ground; and from killing off of steppe rodents as a result of tularemia and other epizootics (Sludskii, 1953).
In years of food shortage, the emaciated animals undoubtedly

die also during flareups of [disease] invasion and epizootics. In

Kolyma

territory,

death of young ermine occurs during a cold spring,

with abundant precipitation and high water in rivers (Belyk, 1958).

The same

is

observed

in the

Volga area
Its

(V.I. Tikhvinskii).

Harvesting has been shown as a very powerful effect on the

process of mortality in populations.
influence
is

selective: the

young, less cautious animals are caught

in the first series,

and then

males

more

often than females.

The

results of selective harvest are

determined by winter food

conditions.

enter traps,

With abundant food, ermine are numerous, but rarely and on the contrary, they go into traps well and are
little

caught in years of starvation with

in

food. Hence, the increase

prepared skins

is in

the season following a

maximum

rise in the

number of ermine, when

there

is

insufficient

food for the

1027
reproducing animals. This usually occurs with depression in number
of mouse-like rodents (Severtsov, 1941; Teplov, 1952).
All carnivorous animals and birds which mainly feed on mouselike rodents

may be

competitors of the ermine.

is

Population dynamics. The ermine population

subjected to

considerable changes. Within the boundaries of the range, these

changes do not proceed equally and simultaneously everywhere,

but
all

the

same

are covering a considerable extent. Within their

borders, subsequent changes in ermine populations are determined

by one and the same similarly directed changes

the external environment.
In the process of these changes,

in conditions of

within different limits: in

Murmansk

numbers of ermine may change district, up to 30-fold

54-fold (Aspisov and Popov,

(Nasimovich, 1948), in the Tatar

observation) by

ASSR

1939), and in the southwestern part of Kalinin district (20 years of

more than 189-fold (Yurgenson, 1959).

In the first

were only 2-6-fold; in the post-war years, the population decreased very sharply, as was also
8 years of observation, fluctuations here

observed

in other districts

of the

USSR.

In northern Kazakhstan,

fluctuations in prepared skins

was 3-7 times, and the catch of individual hunters fluctuated from 2-3 up to 100 individuals; i.e. within 30-50 times (Sludskii, 1953).
Beside the comparatively short-term fluctuations with a time

685

3-4 years between two peaks in ermine numbers in 5-6 years in the southwestern part of Kalinin district and in the upper Pechora 6 years etc., there are, apparently, more prolonged fluctuations within which short-term fluctuations are embedded. Concerning these prolonged fluctuations data are still insufficient. In northern Kazakhstan, periods of depression in numbers last for 5-6 years, and the periods of growth are up to 3-4 years (Sludskii, 1953). The ermine population changes in the course of a year. The greatest number of animals is reached in spring and the beginning of summer, and the least by the end of the harvest season. How great is the significance of the harvest is obvious from the fact that in the floodland of the Kama, the density of ermine at the beginning of the harvest was 52.0 per 10 km^ and by its end only 18.5
interval of

Murmansk

district,

(Zharkov, 1941).

The reasons
1)

for the changes in

changes

in availability

of

its

2) helminth invasion (chiefly in

numbers may be: main foods (voles, water voles); the form of skryabingulosis) and

1028
insufficiently

exposed epizootics;

3) fluctuations in levels of spring

flooding in inundated sections, which with high levels causes the

death of

litters.

The significance of flooding

unsubmerged

is

thus of

two kinds:

moderate floods are favorable, causing a considerable concentration of mouse-like rodents in

parts in the period

is

when
fires,

young are being

fed, but

deep flooding

fatal; 4)

steppe

extensive frozen ground, drying out of steppe water bodies and

spring flooding of the steppe by snow-melts cold rains etc.

Field characteristics. In the snowless period of the year, the

ermine,
its

tracks or excrement are not easily detected. In the period

of snow cover, traces of ermine activity are easily discovered on

the

The ermine moves almost exclusively in jumps, snow paired, sometimes, somewhat obliquely situated, footprints. Tracks and jumps of the males are always larger than those of females, but the tracks of young males may be con-

snow

surface.

leaving on the

fused with those of adult females.

685

Fig.

246. Tracks of large ermine on loose

snow

lying on a crusted surface. Gait

the right, the

unhurried

the prints are situated as a quartet.

tracks.

jumps with paired

scheme of ordinary Neighborhood of Gorkii. 30 December 1916. Sketch by A.N. Formozov, about 2/3 of nat. size.

On


1029

The
mines,

tracks of ermine differ from those of weasel

by

their

larger size, a

much larger search route and by the when walking, sometimes "trots"; i.e. leaves

fact that er-

a triple print

track as in hare or squirrel, which

is not observed in weasel. The ermine digs under the snow significantly more rarely than the weasel, and exits onto the surface after 1-1.5 m. The length of jump of the ermine on the snow varies from 30-

82-100 cm; in weasels their length is 25-36 cm, maximally45 cm. On packed snow, ermine are able to make jumps up to 1.5 m and more (Formozov, 1952), The average length of prints of the male track is 62 mm, of female 46 mm (Nasimovich, 1948). The length of jump of the Altai ermine is equal to 25-50 mm; it sinks into the snow 2-5 cm, and its weight load on 1 cm^ of track is 10 gm (that of sable and Siberian weasel is 12-14 gm, of weasel 7-8 gm) (Dul'keit, 1956). Excrement of ermine takes the form of a spirally-rolled plait thinned at both ends, and consists of underfur and bone fragments. Its thickness is usually about 0.5 cm, and length is not more than
40
to

10

cm

(P.Yu.).

Practical Significance
686

The ermine

is

one of the fundamental items of mass commercial

Its

hunting of our country.

catch in the
this,

USSR

constitutes not less

than half of the world catch. Beside

our country offers ermine

skins of the highest grade to the world fur market (Berezovskii,

The best grade of the North American ermine may only be compared with the 9th grade in
Ishimskii and other standard fur types).
quality type of our standard.

Ermine
its

fur

is

exclusively used for decoration. Because of

this,
is

demand and

price depend, to a great degree, on style

its

and

distinguished by

significant instability.

Regions where ermine harvest has grown up as a specialty are

relatively few.
It is

significantly

more often trapped

incidentally,
is,

or near the vicinity of villages. Therefore, the ermine

All methods for capturing ermine

on the

whole, exploited around our country insufficiently and unequally.

may be

divided into two

groups: hunting with dogs, with or without guns; and capturing

ermine with box-traps and jaw-traps. Hunting with guns cannot be recommended because of damage caused to the skin by shot.


1030
staining the flesh side of the skin with blood, and almost unavoid-

ably dirtying
staining
is

its

fur,

also unavoidable

which lowers the price and quality; bloodwhen an ermine is caught and throttled

by

a dog.

Of
elastic

the passive

means of snaring, metal-arch snap-traps,

kulemka*, and box traps actuated by strong are successfully employed. Less widely dis-

dead-fall

box

traps,

cherkan*

tributed are various nooses (most often, of hair)

which are some"little stool").

times mounted on a frame or crosspiece (called a

All these traps are used as "entrance-gates", on tracks or at holes

and with bait (P.Yu.).

SOLONGOI"
Mustela (Mustela) altaica Pallas, 1811
1811. Mustela altaica. Pallas. Zoographia Rosso-Asiatica,
Altai.
3, p. 98.;

1823. Putorius alpinus Gebler. Memoires Soc. Imp. Natur. Moscou,

6, p.

212. Ridder mines, Altai.

J.

1857. Mustela temon. Hodgson.

Asiat. Soc. Bengal, 26, p. 207.

Sikkim, Himalayas.
1870. Putorius astutus. Milne-Edwards. Nouv. Arch. Mus. Hist.
Nat. Paris, 7, Bull., p. 72.

Mupin
J.

in

Sichuan, China.
Nat. Hist.

1911. Mustela longstajfi. Wroughton.

Bombay

Soc,
566.

20, p. 931. Teza, upper Sutlej valley, Himalayas.

1914. Mustela sacana. Thomas. Ann. Mag. Nat. Hist., 13,

Vic. of Przheval'sk, Tien Shan.

p.

1928. Kolonocus alpinus raddei. Ognev.

Memuary
i

Zool. otdeleniya

etnogr., 2, p. 9. Obshch. lyubit-estestvozn., antrop. Kulusutaev post near Zun-Torei Lake (Tarei-nor), southeast-

ern Transbaikaliya.

1928. Kolonocus alpinus birulai. Ognev. Ibidem,

West Pamir (northern

river) (V.H.).

part of

p. 10. Lyangar in Darvazsk range, Obikhangou

*Local names of traps

^'The
In

is

Sci. Ed.
is

name

is

derived from the Mongolian solongo, and

also

current in Transbaikal.

some

places, the animal

known

as "suslennik"
is

and "mountain polecat". In

the fur trade, the

name "mountain kolonok"

employed.

1031

Diagnosis
Color from bright ocher-reddish to straw-ocher or from nut-brownish to brown. Lighter below than above, transition from dorsal color to ventral color lower part gradual or border sharp. Tail monotone, without black terminal half. Dark coffee-brown field
("mask") on head absent. White field with blurred border on upper and lower lips and chin. Skull relatively elongated (mastoid width
usually less than half condylobasal length of skull); postorbital

687

constriction with abrupt isthmus almost immediately behind

supraorbital processes,
relatively small
its

lateral outlines not parallel.

Dimensions

condylobasal

length of male skull less than 55

(V.H.).

mm,

of female, less than 48

mm

Description
In general appearance, dimensions and habits, the solongoi
is

very

similar to ermine, but differs in

its

somewhat
Tail length
1/2.

larger head, longer

and

fluffier fur, the

more

furry

tail.

more than

1/3

body
on

Claws white. about with terminal Winter dense, and although short (length of guard more than ermine), quite sacrum 12-18 mm only
length
hairs,

fur

hairs

slightly

luxuri-

ant. Tail
its

covered by hairs of equal length and equal thickness over

as in ermine, but

whole length
tail

somewhat more

luxuriant. In

some forms,
riant.

hairs very strongly lengthened,

even more luxu-

Soles of feet completely covered with hairs; digital and sole

pads not visible.

In winter fur (Altai), general color tone of upper side of

body

pale straw-ocherous or clay-ocherous; guard hairs are shiny, with

brown
head
is

tips. In

middle of back, color more intense. Upper part of

duller than back (here yellow tone of
"little

somewhat darker and

underfur drops out), but

cap" very weakly defined in form

at all.

of darkish overlay, sometimes not developed

Dorsal color of

sides gradually passes into considerably lighter straw-ocherous with

yellowish-red tones or to whitish-ocherous color of venter, chest

and throat. Chin, most anterior part of lower surface of neck, lower and upper lips whitish or white or almost white. White color of lower neck surface imperceptibly passes into color of lower part of body. Concerning white areas on upper lips, the same applies, but
these areas are usually quite sharply demarcated from color of

1032

upper head.

On

throat, chest,

and base of paws, vague whitish

spots sometimes occur.

On

neck, dorsal, as well as lateral color gradually passes into

ventral color; on head, borders of colored areas quite sharp

due

to

described light fields. Small dark spot on light background usually

dish
ter.

found behind corner of mouth. Tail same color as back, with redtint, similarly colored above and below, i.e. darker than ven-

Upper

parts of legs

same color

as back,

and lower parts

as

sides, inner parts colored like venter. Feet dorsally lighter than

back

straw-colored
it

or whitish.

Covered ventrally by dense whitcallosities.

ish-silvery hairs

which conceal

Summer

fur (Altai) considerably shorter, coarser

tail

and more

sparser than winter fur. Hairs on

sparser, causing

considerably shorter and

surface of

to appear thinner.

Lower

paws more

688

weakly furred, digital and sole pads quite noticeable. Color of summer fur very sharply different from that of winter and much darker than it. Upper side of body dark brownish-ocherous color, more intense in middle of back. Sometimes color of spine weakly delineated against general background of back. Dark area of middle back extends along upper neck and covers top of head. Sometimes top of head somewhat darker than middle back, and darker "cap" without sharp outlines marks itself on head. Sides somewhat lighter than back nut-brown. Lower body lightocherous, even whitish-ocherous, throat slightly brighter. Border between color of lower body and the sides completely sharp,

almost as in

summer

ermine, but situated

much

lower.

Lower
sides, fur
in

surface of

paws dark,

like back,

upper has color of

1

on

digits whitish above.

Sometimes, small (about

cm

diameter) dark spots of same color as sides occur on lower

surface of body. Region between rami of lower jaw lighter than

general color of lower body, sometimes whitish or almost white.

Upper and lower

lips, a

lips

white or whitish. This color on lower jaw

field usually quite sharply

imperceptibly passes into color of chin, throat and neck.

narrow white

On upper demarcated from

lips

dark color of head. Sometimes, white color of upper

poorly

developed

narrow or weakly lightened and gradually merges with

is

dorsal color. Behind corner of mouth, a bit below border between

top and bottom, a small chestnut dot occurs.

Individual variation in color

usually of a fluctuating type

in color is

and of small amplitude. Sexual dimorphism

absent.

1033

Young up

to

two months are covered by a

is

short, fluffy, dull,

dark

fur. In the first winter,

they are clothed with adult pelage. Geoquite significant and

is

graphic variation in color

expressed in

general tone of winter and

summer

fur

either lighter or darker

depending on brightness, degree of contrast between winter and summer fur, sharpness of border between dorsal and ventral color, and in presence or absence of border itself (gradual transition)^'*, intensity of ventral color, which may be almost white and other
characters. Essentially, this variability, in this way, bears not only

a quantitative but also a qualitative character.

The

skull of the solongoi

is,

in

some

respects, similar to that

it

of the Siberian weasel [M. sibirica], but differs from of essential characters.
It

in a series

is

not so narrow, long and elongated,

having a shorter, wider and more swollen braincase. The region of the postorbital constriction is relatively short with a well defined
isthmus, located near the supraorbital processes.

The narrowest

part of the postorbital region ("isthmus") lies on the line connect-

ing the supraorbital processes, at a distance less than half the

distance between the ends of these processes.

The

lateral outlines

of the postorbital region of the skull (region of the frontal bones)

are not parallel (or

more or

less parallel), but

form angles, the

apices of which are directed towards each other (toward the sagittal

plane of the skull).

Infraorbital foramina small, oval, their longitudinal diameter

less

than or equal to longitudinal diameter of upper canine

its

alveolus. Interpterygoid area usually narrows forwards, and

general outline has an angular form, apex directed forwards. Auditory bullae are elongated, in plan a
^This distinction

smooth rectangular form,

noted nor emphasized in our

between the color of the summer and winter fur is neither literature, first of all; and then, that in the summer solongoi there may be such an "erminish", bicolored type. Usually, even in the time of Satunin and to this day guides and monographs published in 1962 and 1965 (edited by I.I. Sokolov, A.N. Formozov and A. P. Kuzyakin) have on the contrary underlined, contrary to other groups of the genus, that in opposition to other groups of the genus "color of the dorsal and ventral sides ... is monotone-yellowish-red" or "the color ... on the back and on the belly is the same", "color of the back and abdomen is identical... "color of dorsal and ventral sides of the body is red, dark-yellow or grayish yellow" etc., and that the solongoi is completely identical to the Siberian weasel in color. Because of this, color is considered one of the main characters for the group "Kolonocus" and is considered as such in keys, diagnosis etc. So categorical a formula not only
presents an incorrect representation of the species characteristics, position and geographic variation, but also
its

systematic

makes

difficult the

simple identifi-

cation of the solongoi in several regions.

1034

689

Fig. 247. Skull of solongoi

Mustela (Mustela) altaica

Pall.

1035
with inner borders of almost parallel lines. Width of auditory bullae

approximately equal to half

690 interpterygoid depression.

its

length. Distance

between bullae

usually greater than width of the hard palate at anterior part of

Zygomatic width approximately correin mastoid region. Zygomatic arches weak. Protuberances, crests, etc. unnoticeable; sagittal crest weakly
sponds
to

width of skull

defined, occipital crest

somewhat

larger.

Frontal area elevated,

it

profile of facial region slopes

down from

quite suddenly. Teeth

relatively strong.

On

the whole, the solongoi skull

it

by comparison with the

its

Si-

berian weasel skull with which

infantile features.

is

usually affiliated, has several

As regards degree of

development,

it

may be

equated with the ermine skull.

Age and

sexual differences in the solongoi skull have not been

specially described. In general, they correspond to those

men-

The female skull is considerably smaller and somewhat lighter than the male skull, with less defined protuberances, crests, etc. Skulls of young animals have more brain case, which is well marked in animals up to one year old,
tioned above for ermine.

but in the second winter of

life,

the skull acquires

all

the features
is

of the adult animal. Individual variability in the skull

great, but

does not exceed that

in

other species. Geographic variation of the

skull, at least within the

boundaries of the

USSR,

is

neither devel-

oped nor noticeable, with the exception of one

Structure of the os penis
is

race.

typical for the majority of the

species of this genus,

i.e.

distal

end forms an upwardly bent hook.

Dimensions of the solongoi are close to those of ermine. Body is 224-287 mm, of females, 217-249 mm; tail length of males is 108-145 mm, of females, 90-117 mm; length of hind foot in males is 39-47 mm, of females, 33-45 mm; height
length of males

of ear in males

is

16-21.6

mm,

of females, 14.0-20.2

mm (Sludskii,

1953; Stroganov, 1962).

Condylobasal length of male skull (14) is 46.3-52.8 mm, of females (17), 40.0-47.1 mm; zygomatic width of males is 23.127.6 mm, of females, 19.0-24.2 mm; interorbital width of males
is

8.8-12.2

is

mm, of females, 7.7-9.5 mm; mastoid width of males 20.2-24.0 mm, of females, 17.7-22.2 mm, (Stroganov, 1962).
Length of OS penis
is

25-27.1

mm

(Ognev, 1931).

1036

Weight of Siberian solongoi males is up to 350 gm, of females up to 220 gm (Stroganov, 1962), usually much smaller. Males from Pribalkhash weigh 217-255 gm (January-March), females, 122-135 gm (Sludskii, 1953) (V.H.).
Systematic Position

Due

to the

known complexity and

diversity of structure of genus

Mustela, determination of the position of separate species presents

difficulties.

Of

those forms of the genus which are accepted at the

present time as independent species, M. altaica stands closest of

all to

the southern Asiatic (South China, Indochina, Himalayas)

It is

M.

kathiah.

sometimes considered, and is possibly true, that it is only a race of M. altaica. Relationship to the Malayan M. nudipes
insufficiently clear, but they are probably close.

is

Among

our forms and in our literature, M. altaica

is

accepted,

according to a tradition proceeding from Satunin, to be closest to

emphasized by combining one genus) Kolonocus. In keys for identification, both species, even in the most recent books, are found in one group (see note on page 1033). Nevertheless, in actuality there is no particular relationship between the solongoi and Siberian weasel. The singular, or in all events the main common characteristic of both species was considered (as in keys) to be color. However, there are very essential
the Siberian weasel,
sibirica. This is
(in

M.

them

into

one subgenus

extreme cases

differences in this characteristic.

As shown

(see above, section,

"Description", and below, "Geographic Variation"), in the solongoi,

color type itself

is

different, particularly contrast

between dorsum

and venter may be very strong, and the demarcation between one and the other is sharp. The general color tone in some forms [of
altaica]
is

almost devoid of the characteristic ocherous-reddish

is

tone. External similarity to the Siberian weasel

found

in

only

691

one race of the species (M. At the present time, it

solongoi
is,

a.
is

raddei) and only in winter pelage.

more correct

to consider that the

according to several external characteristics and crathat as a

It is necessary, however, consequence of the significance of some

niological peculiarities, closer to ermine.

to

have

in

view

race characteristics of our solongoi, not

all

of them are identical

with them in this connection. Thus, M.

a.

raddei possesses

all

known

characteristic similarities of the Siberian weasel, not only

1037
in color but also in the craniological structure.

With

all

of that, the

solongoi

is,

in all events, closer to

ermine than

to Siberian weasel.

According to the sense of some (Pocock, 1941), differences between solongoi and Siberian weasel are even larger than those between Siberian weasel and European mink (M. lutreola). This
recital lead us to consider the order of species

accepted here,

i.e.

the structure of the genus, as a

and refuse
in

to accept the

more suitable natural relationship group Kolonocus in any form or rank, or

general any preference for affinity of solongoi and Siberian

weasel within the genus Mustela (V.H.).

Geographic Distribution

The range of

the species

is

relatively small

rarely level regions of Central Asia

and occupies montane, and adjacent parts of Middle

Asia and Siberia, the Far East and China.

Geographic Range
In the

in the Soviet

Union

it is small, and represents the most northeastern, fringe of the species range. It consists and northwestern northern of several, in part very small, sections separated from each other

USSR

within our country but united to the south, outside the border of
the

USSR. The

details

solongoi distribution are

and even some of the essential features of still poorly known and the range cannot
occupying both

be outlined with sufficient accuracy.

In the west, the solongoi inhabits the Pamir,

eastern and western parts of the region and the Trans-Alaisk and,

apparently, the Alaisk range.

The northwesternmost known point

Lyangar
in the

of occurrence in the Pamir

is

upper Obikhonogoi

(Vakhsh)

in the northern part

of the Darvaz range. In southern

Tadzhikistan, in turgais along rivers flowing into the Pyandzh and

Amu-Dar'ya,

it

is

absent. Concerning

its

occurrence

in the Peter

the Great, Gissar and Zeravshan ranges, there are no direct refer-

ences. At least in the latter two regions and along their western
spurs,
it

is

absent (in general not recorded within Uzbekistan;

It is

Ishunin, 1961).
In the

also, apparently, absent in the

Turkestan range.

montane system of Tien Shan, the solongoi lives in its eastern part in Kungei Alatau, Tersk Alatau and in Zailinsk Alatau, and in montane regions lying to the south as far as the Fergana


1038
valley.
In this valley the solongoi has not

been noted; and

is,

apparently, absent there. The Tien Shan and Pamir sections of the range, are probably in the extreme eastern, montane parts of Fergana,

but within the borders of the

In the

USSR may be isolated from each other. mountains of u'estern Tien Shan (Kirghiz and Chatkal ranges, Talass Alatau), presence of the solongoi has not yet been recorded and it probably is absent there (Sludskii, 1953); mention
of
its

sporadic occurrence "all over Kirghizia" (D. Dement'ev,

1938) and several others are too indefinite. It is encountered in the Dzhungarsk Alatau, in the lower Hi and Karatal, along the Lepsa and Aksu, along the southern shore of [Lake] Balkhash (in the east, recorded at mouth of Ayaguz) and in the Alakul' depression. Here, the occurrence of this montane species among tugais and
reeds on the plain between deserts
Farther to the east,
it

is

remarkable.

lives in the Tarbagatai, Saur, in Zaisan

depression and the mountains surrounding

Irtysh and in the Kalbinsk Altai
(left

it, along the Chernyi bank of Irtysh). In the Altai, the solongoi is known from a series of places Sailyugem, Chuisk mountains and Chuisk steppe (Kosh-Agach) in the south, from the Ust'-Koksa, Uimon, Ongudai, Koop-Ching, Kotanda Elikmonar regions, and a series of other places, however, not to the east of the Katun'. The northern- and northeastern-most known point of inhabitation in the Altai Turochak lies, how-

693 ever,

on the Biya (northeast of Gorno-Altaisk); reference to

mountains south of
it.

its
it-

occurrence around Barnaul applies, apparently, not to the city

self but to the Altai

To

the west of the Katun' the solongoi lives, apparently,

it is

all

met with in the south up to the frontier (Kurchumsk, Narymsk and other ranges), and in the north around Ust'-Kamenogorsk and extends to Semipalatinsk and it even penetrates somewhat farther north
over the mountains and foothills as far as the Irtysh;
(Beskaragaisk region of Pavlodarsk
region of occurrence
is

district). In this

way, the Altai

united with that of Turkestan in our country.

Deep
.

in the eastern part of the Altai to the east

is,

from the Katun',

the solongoi
foothills).

apparently, absent (Turochak lies in the northern

it

Thus,

has not been discovered in the Altai preserve

around Teletskoe Lake and along the right bank of the Chulyshman (Shapshal range; V.G. Heptner, Yurgenson, 1938). It is also absent along the West Sayan range and the mountains to the north of it,
as well as in Tuva.

'a
)
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1040
In the literature, very general references to the occurrence of

the solongoi "in Altai and in

and sometimes,
emphasized. As
record
is

its

Sayan" are occasionally encountered, occurrence in all of these montane regions is

is

is

apparent, in the Altai the picture

complicated,

and as regards the western Sayan, not a single accurate positive

found, but there
is

negative (Stroganov, 1962, after

in this district with a question

Nadeev). There are also no positive data concerning Tuva

(Yanushevich, 1952, refers
the Trans-Baikal solongoi).
to
it

mark; see also the section "Geographic Variation" and notes on

The Cis-Baikal
Baikal
is

part of the range of the solongoi* west of [Lake]

represented by a quite narrow strip which, beginning from Achinsk and Irbeisk regions (east of Krasnoyarsk, about 95" E long, and 56 N. lat.), extends along eastern Sayan range to the
southeastern corner of Baikal, including in particular, the Tunkinsk

mountains and Slyudyank region on the east (southeastern extremity of Baikal); it continues onto Khamar-Daban (Podarevskii, 1936). Thence to the south, the range goes onto the boundary with the Mongolian People's Republic (Prikosogol'e).

If all existing

information about the absence of solongoi in the

western Sayan, in Tuva and in the extreme eastern parts of Altai, the Eastern Sayan, Cisare true then, it is evident that these

Baikal and Trans-Baikal areas (see below) of the species distribution are cut off, within the borders of the

USSR, from

the Tien

Shan-Altai part. At the same time, the reasons for this probable gap are not known, and the occurrence of the solongoi in western

Sayan remains likely. However, the well manifested differences between the races of Altai and Trans-Baikal solongoi speaks against this (the eastern Sayan [race] is unknown; see below, section "Geographic Variation"). In Trans-Baikaliya, the solongoi is widely distributed. The eastern limit of the range here to a significant extent is served by
the [eastern] shore of [Lake] Baikal.

Southward the range extends

it

beyond the

state frontier,

except that

also occupies the steppe

regions of both southwestern and southeastern Trans-Baikaliya (for

example, Aginsk steppe, the steppe expanses of Olovyannin region

etc.).

The

distribution of solongoi to the north

beyond Baikal

is

not well elucidated. Apparently, the range goes to

the Lena, and

In Russian original, "kolonok," a lapsus

Sci. Ed.

1041
lat. and somewhat to by the species). Thus, the solongoi is known (V.D. Shamykin) from the area along the Lena between approximately 111-112'' and 120 E. long. The border of the range in the northwest between the northern extremity of Baikal and the Lena, is unknown.

perhaps somewhat beyond

it,

i.e.

to 60**

N.

the north (the northernmost place inhabited

The

distribution to the east

is

is

also not accurately established.

In the northeast, the solongoi

known from

the
its

Aldan (Tugarinov,
upper reaches, and
confluence of

Smirnov and Ivanov, 1934), apparently from

in the

south

it

undoubtedly reaches
is

in the east the

the Shilka with the Argun' and

the
694

recorded in the western parts of

1936)".

Amur

district (Plyater-Plokhotskii,

For the distribution

to the east of the

meridian of the conflu-

ence of Shilka, Argun' and Aldan the literature contains only two positive indications in the western part of Amur district and the

neighborhood of Blagoveshchensk (Stroganov, 1962). In several works dedicated to the regions north of the Amur, this species is
and is usually considered to be absent there. In actuality, the range apparently covers the whole expanse between the Amur and the Stanovoi range, perhaps without reaching it, but, occupying the upper reaches (and the sources) of the Zeya and Selemdzhya. From here, it descends abruptly to the south,
not referred to at
all

695

occupying the lower Bureya and the southwesternmost (southern) part of the Bureinsk range^^.
Including in this

way

the eastern slopes of the southern ex-

tremities of the Bureinsk range^^

693
^'In

somewhere

little

to the

west of

the description of the solongoi range east of Baikal, besides the sources

S. Anaskin (Bur[yat] ASSR), P. Zimin 694 (Chita) and D. Ivanov (Yakutsk) were used. These available data are contradictory. Thus, for Chitinsk district, showing the solongoi distributed in the Krainii (Extreme) North in Yakutiya, its presence is denied. For Buryat ASSR, the northern border of the range is given as follows: from the eastern shore of Baikal at a place about 100 km. south of the mouth of upper Angara, to the source of the Barguzin, then to Baunt Lake and from it to the northeast, at first slightly to the south of the Mui river and to the Vitim to its mouth. Therefore, the northern and eastern borders of the outlined section of the range is preliminary and to a certain degree, conditional. They strongly differ from those advocated by other authors, notably Stroganov (1962). ^*The above described part of the range in the Priamur'e is given according to range maps made by A. Samsonov (Blagoveshchensk) and V.P. Sysoev (Khabarovsk). "Range in Primor'e after data (map) of P. Bel'skii and V.D. Shamykin. The

mentioned, unpublished information offered by

solongoi

is

very rare everywhere. In the faunistic literature concerning the territory,

is

the solongoi
fact

more

restricted than

nowhere mentioned. Therefore, was mentioned.

it is

possible that the range here

is

in

1042
30
60

694

Fig. 249. Species range of the solongoi,

Mustela (Mustela) altaica

Pall.

(V.G. Heptner).

132 E. long, or along

exits into China.
at
It

it,

the range border crosses the

again enters the borders of the

Amur and USSR in Primor'e,

the Bikin Basin in Ussuri. Thence, the range extends as a

strip

narrow

southwards along the Ussuri valley and the adjacent

bit

western foothills of the Sikhote-Alin', and then, a

latitude of the southern extremity of slightly to the

south of the

Lake Khanka, southeast and passes to the Ocean

the border turns

at

Nakhodka.
in the

Within the borders of our country, a clear break

is

range

marked out along the Amur to the east of the Bureinsk range and along the lower Ussuri. However, it is not excluded that, in the

1043
Priamur'e, the solongoi goes to the east somewhat farther than

mentioned. ^^

Geographic Range outside the Soviet Union

This
is

only

known

in

an entirely general outline.

It

occupies

Kashgariya (obviously, montane parts contiguous

to our country)

and Dzhungariya, in the Mongolian Republic Mongolian Altai and the eastern part of the Mongolian Republic from the Kentei to the Great Khingan [mountains] and south approximately to 46 N. lat. (in the region of the Great Western and Gobi lakes and in the

Khangai
solongoi

as well as in the remaining parts of the country, the

is

absent). In the

Korean Peninsula,

it

inhabits at least

its

northern parts. In China, the range occupies

parently, to the Great Chinese Plain,

its

northeastern part,

extends eastwards to the middle parts of Shansi and to Hubei (apcorded), and southwards to Sichuan,

from whence it Yunnan and Fujian,

is

not re-

as well as

Tibet. In the south, the range includes the

to

Himalayas from Sikkim

Kashmir, Ladakh and Gilgit, the Karakorum and the extreme

east of Afghanistan (V.H.).

Geographic Variation
Within the borders of our country, the solongoi provides several
well defined races.
territory
It is

noteworthy that despite the relatively small

is

occupied in our country by the range of the solongoi, the

these races
quite large, and morphological differ-

number of

ences, at least some, are significant.

They

are greater than in races

of our ermine, even those inhabiting parts of the country that are

remote from each other. The subspecies of solongoi represent also

interesting combinations of characters together forming not only

quantitative but also

city

some

qualitative differences.

Due

to the scar-

of material in our museums, geographic variation of the

^Range in the

USSR

after

1870; Gassovskii, 1927; Emel'yanov, 1927; Kuznetsov, 1929,

Maak, 1859; Shrenk, 1859; Radd, 1862; Przheval'skii, 1948, 1950; Ognev,

1931; Tugarinov, Smirnov and Ivanov, 1934; Flerov, 1935; Zolotarev, 1936; PlyaterPlokhotskii, 1936; Podarevskii, 1936; Favorskii, 1936; D. Dement'ev, 1938; Yurgenson,

1938; Kolosov, 1939; Razoprenova, 1939; Bobrinskii, 1944; Kopylov, 1948; Nasimovich, 1949; Yanushevich, 1952; Sludskii, 1953; Chernyshev, 1958; Afanas'ev, 1960; Ishunin, 1961; Stroganov, 1962 and other sources and according to unpublished material of S. Anashkin, D. Ivanov, G.F. Bromlei and V.G. Heptner.

1044

however, quite insufficiently studied. Within the USSR, the following forms now can be recognized from material of the Z[oological] M[useum] of M[oscow] U[niversity]; measuresolongoi
is still,

ments of the Siberian forms are mainly from Stroganov, 1962. 1. Turkestan solongoi, M. (M.) a. sacana Thomas, 1914 (syn.
birulai).

General color tone of upper body in the winter fur very light, straw-sandy-ocherous. Venter considerably lighter than dorsum,
696

from color of back to color of belly gradual. fur, the color above is noteably darker than in winter, brownish-gray with light-chestnut and reddish tones. Bases of hair dorsum bluish-gray. Top of head darker than back (pale-sandy and
transition

In

summer

smoky
away
very

tints

of underfur not present). Pure white, or with light

tints

straw-colored

on

lips,

chin and upper part of throat washed

at edges, rarely sharply demarcated from color of throat and

cheeks. Color of lower surface of the body (belly, chest and neck)
light,

pale-straw or ocherous-white, in

some animals almost

white (bases of hairs often white) with yellowish or reddish overlay, particularly obvious on sides of neck. Dark color of dorsum
extends to sides very sharply demarcated from light color of venter. Distal portion of feet silvery- white above. Tail moderately fluffy.

On

the whole,

summer

color

is

variable, but lighter than in other

races (material from Issyk-Kul').

224-282 mm, of females, 218-230 116-132 mm, of females, 90-114 mm; length of hind foot of males is 39-46 mm, of females, 34-35 mm; ear height of males is 16-20 mm. Weight of males (winter) is 217-255 g, of females, 122-135 g

Body
tail

length of males

is

mm;

length of males

is

(lower Hi, Pri-Balkhash; Sludskii, 1953).

Mountains of Middle Asia including Dzhungarsk Alatau,

Balkhash.
Outside the

Pri-

USSR probably

in

contiguous parts of Kashgariya,

perhaps in Dzhungariya and Afghanistan.

Altau), the lightest form within the

form (material from Tersk both in summer and in contrast between dorfur and winter. In general tone of summer sum and venter of the body it is very reminiscent of ermine. Nomenclature of this form is not completely clear. It is undoubtedly (contrary to Ognev, 1931) well differentiated from the Altai [form]. The identity of the Tien Shan and Pamir solongoi
It

is

a very sharply distinguished

USSR

1045
("biruleV)

is

highly probable, but needs confirmation.

On

the other

hand,

it is

probable that the Pamir and perhaps also the Tien Shan

are related to the

Himalayan form, temon. Their description

the
is

is

similar and there are analogies to such distribution found

among
to

other

mammals. Therefore,

same sacana given

provisional.

the

Turkestanian montane solongoi

The systematic position of solongoi of

Pribalkhash'e
2.
is

the river valleys in

is

not studied, but

its

position in this race

sufficiently evident.

Altai solongoi

M. (M.)

a.

altaica Pallas, 1811 (syn. alpina).

in

Color of winter fur of dorsum darker than

light ocherous-chestnut color, dull.

preceding form,

Lower

surface of body

much

between color of dorsum and venter gradual, but in some, it is demarcated by more or less distinct boundary. Tail is moderately fluffy, color as on back. In summer fur, dorsal side dull and very dark-brown tone. Darkening along middle of back relatively stronger than in other forms. Venter considerably lighter than dorsum, dirty light-ocherous tone, sometimes with small brownish spots. Border between color of dorsum and venter sharp. Body length of males (7), 235-M241-260 mm, of females (5), 217-M223-227 mm; tail length of males, UO-Ml 29-145 mm, of
lighter than dorsal. Transition

104-M108-113 mm; length of hind foot of males, 41.0M44. 1-47.0 mm, of females, 35.5-M37.2-38.4 mm; ear height of males, 20-M20.8-21.6 mm, of females, 15.7-M16.8-18.2 mm. Condylobasal length of male skull (7), 51.2-M52-53.0 mm, of
females,

females

(5),

46.8-M47.3-49.5

mm;

zygomatic width of males, 26.2-

M27.2-28.4 mm, of females, 23.2-M24.4-25.7 mm; interorbital width of males, 10.3-Ml 1.3-12.6 mm, of females, 9.3-M 9.6-10.2 mm; mastoid width of males, 22. 8-M23. 3-24.0 mm; of females, 22.0-M22.4-23.2 mm.
697

In Altai.

Outside the
golian Altai).

USSR western

part of

Mongolian Republic (Mon-

A
3.

well characterized form, obviously distinguished from both

eastern and western forms. Darkest form in

USSR.

Trans-Baikal solongoi M. (M.)

a.

raddei Ognev, 1928.

In winter fur, dorsal color bright reddish-ocherous or pale-

ocherous. Abdominal surface of body slightly lighter than back.

1046

from back color to abdominal color gradual. Tail bright similar to back and fluffier than other forms. In summer fur, color of dorsum dark-chestnut, but somewhat lighter than in Altai form (altaica); venter lighter than dorsum ocherous-yellow, of quite intensive tone. Transition from color of dorsum to abdominal gradual, without sharp border. Skull dimensions less than in forms altaica and sacana, somewhat more elongated and sturdy, with narrower interzygomatic
but also light-ocherous.
sides, transition

On

region.

Body length of males (6), 249-M265-280 mm, of females (7), 222-M224-226 mm; tail length of males, 108-M129-144 mm, of females, 100-M106-117 mm; length of hind foot of males, 40.8M42.3-44.2 mm, of females, 33.2-M36.9-43.8 mm; ear height of males, 17.4-Ml 8.3-26.6 mm, of females, 16.0-M17.3-20.2 mm. Condylobasal length of male skull (6), 49.5-M50.5-52.3 mm, of females (7), 43.0-M44.2-46.2 mm; zygomatic width of males, 25.7-M 26.6-27.3 mm, of females, 20.0-M22.6-24.5 mm; interorbital width of males, 9.3-M10.3-11.8 mm, of females, 8.1-M9.210.0 mm; mastoid width of males, 20.2-M22.5-23.8 mm, of females, 18.7-M 19.0-1 9.4 mm. In Trans-Baikaliya, southern Yakutiya, Priamury'e? and Ussuri
Territory.

Outside the

USSR,

in eastern part

of M[ongolian] P[eople's]
a very well character-

R[epublic], and northeastern China.

The Trans-Baikal solongoi represents

ized form. In winter pelage,
it

is

the brightest and

most beautiful

solongoi.
Altai and

Summer

pelage

is

a little lighter than in the neighboring

somewhat

similar to the fur of the Tien Shan; absence

(weak development occurs rarely) of a sharp boundary on the sides between color of dorsum and venter is characteristic. Characteristics of solongoi of the western Sayan are not known, nor is the
systematic position of solongoi of Ussuri Territory.

The sharp differences between

parts of the Altai

Altai and Trans-Baikal solongoi

in the eastern

confirm the opinion about the absence of the species

and eastern Sayan. The differences mentioned between the eastern Baikal region (Trans-Baikaliya) and western (Tien Shan and Altai) solongoi and in general the outlines of the northern part of the range (see Fig. 249) make us think that, from the old southern parts of the range, the solongoi dispersed from the


1047
southeast into Trans-Baikaliya, and from the south and southwest
into Turkestan

and the

Altai.

Outside the

USSR,

the form
is

M. (M.)

a.

temon Hodgson, 1857*

(syn. astutus. longstaffi)

usually recognized, from the Himalayas

and Tibet.

Geographic variation
borders of our country
in species

in

solongoi

occurring

outside

the

demand reexamination, and nomenclature

rejected by the majority of authors (V.H.).

revised. Inclusion of "M. kathiah" (with the subspecies caparioccoi)

M. altaica

is

Biology
Population. Within the limits of
distributed and usual, but
its

range, the solongoi

is

a widely

uncommon

animal. They constitute one

of a few, except in the lower reaches of the rivers of southern

Pri-Balkhash, where in years of abundant animals, along a route of

10

km

there were 8-25 solongoi tracks.

Habitat.
698

The most
is

significant part of the range lies in

montane

regions where there

negligible development of forest vegetation

extremely varied. They

and a predominance of open landscapes. Habitats of solongoi are are, first of all, determined by interrelationships with the other small carnivores

by
in

competitors. In

Kazakhstan, competing species reveal themselves to be ermine,

marbled polecat and kolonok. Kolonok and solongoi are two such inter-relating species. In many cases, they occupy similar habitats:
valleys of

montane streams and lakeshores

montane-taiga

re-

gions. In these cases, solongoi are usually restricted to subalpine

and alpine zones, and kolonok to forest zones, the borders of which solongoi rarely penetrate, and then only where kolonok are absent
(Sludskii, 1953).

altitudes

mountains of Middle Asia, the solongoi is typical of from 1400 to 3000 m above sea level, inhabiting the shrubby valleys of montane streams, rock slides and talus covered with herbs, pebbly areas at the upper borders of spruce forests [scree slopes?] and other places (Shnitnikov, 1936; Sludskii, 1953).
In the

*In Russian original, "1957"

Sci. Ed.

'(.;

1048
In Tien

Shan

habitats,

where there are many ermine, there are few

solongoi and vice versa. In the vast areas of channels in the deltas

of the rivers draining into [Lake] Balkash (Sludskii, 1953) which

and sandhills, the most often encountered along the banks of small river channels and lakes, in small meadows with dense herbaceous cover and were rarely in tugai [riparian] forests. Sandhill, here are inhabited by the marbled polecat [Vormela peregusna].
alternate with small lakes, salt flats, ridges

solongoi

is

In western Trans-Baikaliya, the solongoi

is

common

in

steppe

and forest-steppe sections. In upland steppes, the solongoi is characteristic of hollows and ravines with mesophytic vegation (Fetisov,
1937).

The same

applies to places in southeastern Trans-Baikaliya

(P.B. Yurgenson).

The solongoi does not avoid human proximity,

houses and dwellings.
in part, fish constitute the

settling in store-

Food. Rodents and birds of moderate and small size, lizards and main food of solongoi. Cases of attacking

domestic birds and stealing eggs and other food products are known.

Winter foods of the solongoi in western Trans-Baikaliya; 22.7% and insects 3.1% (97 rodents consists of 73.2%, birds spec). Daurian pikas [Ochotona dauricd] occupy first place in

order of frequency (see Table 65).

predominate

The narrow skulled vole, Brandt's vole and Daurian pika in the summer food of solongoi in southeastern Trans-

Baikaliya (Brom, Vovhinskaya and Fedorova, 1948).

Table 65. Seasonal foods of solongoi in western Trans-Baikaliya (% of occurrence)

Type of food
Daurian pika
Small birds
Brandt's vole [Microtus brandti]

Winter
34.2
13.4
9.3 9.3 8.2

Summer*
27.2
0.9

27.2
36.5

Narrow-skulled vole [M. gregalis]

Daurian partridge[Par^A: dauricae] Michnoe's vole [M. fortis michnoi]

6.2
5.1 5.1

Clawed

jird

[Meriones unguiculatus]

Large Asiatic wood mouse [Apodemus

peninsulae]

Shrews
Daurian hamster [Cricetulus barabensis]
Fish

3.1

2.1

9.1

1.0

Summer
steppes).

food data after material from southeastern Trans-Baikaliya (Daurian

1049
699

In Pri-Balkhash'e, the solongoi feeds mainly on voles and

mice

and

in

summer,

to significant

degree on lizards (Sludskii, 1953)

(see Table 66).

Table 66. Seasonal foods of solongoi in Pri-Balkhash (% occurrence)

Type of food

Winter
61.0 61.0 24.2
14.7 10.5 1.0

Summer
66.6 66.6 25.2 13.9
5.5 8.7
11.1

Mammals
Rodents Voles, mice Muskrat Tamarisk jird. [Meriones tamariscinus]

Midday gerbil [M. meridianus] Water vole [Arvicola terrestris] House mouse [Mus musculus]
Tolai hare
Birds

[Lepus tolai]

Pheasant

[Phasianus colchicus]

Moorhen [Gallinula chloropus]

Ducks
Small birds
Lizards

Frogs
Fish
Insects

5.2 2.1 3.1

32.6

5.8

8.3
11.1
2.1

5.5

2.7 2.7 22.5 2.7

8.3

Coagulated blood
Total
data

22.8

95

36

Home

range. Data are absent.

shelters. In the

Burrows and

for rearing young, the solongoi

way of temporary burrows, and more often use holes dug by

rodents (water vole, muskrat, gerbil, pika). Therefore, the burrows

of solongoi are often located near water and even on floating mats

on lakes (Sludskii, 1953). The burrow usually has a nest chamber and also a "latrine". At its entrance, remains of food are often found. In the autumn-winter period, the solongoi has several temporary shelters within its home range in each of which it lives for

some days

in a

row.

Daily activity and behavior. In summer time, the solongoi is active in twilight but is also met with during the day, i.e. its daily

rhythm is imprecise (P.B. Yurgenson Trans-Baikaliya; Sludskii, 1953 Kazakhstan). The solongoi is bold and has little caution. Being a typical terrestrial predator, it can also climb well and does not fear water. Its voice is a shrill chatter like that of ermme.
activity

1050 Seasonal migrations and transgressions. Data are absent. Reproduction. Estrus and mating of solongoi in Kazakhstan
are observed

from the second

half, rarely

from the beginning, of

January to the beginning of April. The duration of pregnancy has been given as from 30 to 40 days. The number of young in a litter is from 1-2 to 7-8. The
latest date of parturition

was noted on 20

May

(Sludskii, 1953).

According
first

to

more

detailed investigations (Gusev, 1955), the

signs of spermatogenesis in individual animals (210 spec.)

were observed at the end of December. In the first half of January, males. Extinguishing of the process it was observed in all extended from the second half of March to the first days of May. In individual old females, estrus appeared in the first half of Febin March. In young, estrus lasted from ruary, and in the majority

the second half of

March

to the first

days of April. Parturition

700 Duration of

proceeded from the first days of April (old females) to 15-20 June. pregnancy ranged from 30-35 days. The number of embryos was from 2 to 6; average for adult females was 5.5 and 3.6. Lactation period lasts 34-41 days. for young

Growth, development, and molt. Eyes of the young are opened after 27 days, probably at the age of 30 days. At the age of about 2 months, they already lead an independent mode of life, but the
does not break up until late autumn (Sludskii, 1953). Enemies, diseases, parasites, mortality, competitors, and population dynamics. Ermine, kolonok and marbled polecat are considered to be the main competitors of solongoi. Significant mortality (cause not established) was observed in 1941 (individual
litter

animals and

litters) in

Semipalatinsk district (Sludskii, 1963).

Fluctuation in numbers of solongoi are well expressed. In

periods of depression, their
that, restored in the

number decreases 2-4 times and

fell

after

course of 2-3 years.

From 1936-1949, num-

bers of solongoi in Kazakhstan

two-fold (Sludskii, 1953). In

1939, the sharp decrease in

its

numbers coincided with a strong

tularemia epizootic

among

rodents.

Numbers of solongoi
is

are also,

without doubt, affected by the vast spring fires in the delta of the
Hi river. In southeastern Trans-Baikaliya, the picture
different.

Here, there were four increases in numbers of solongoi within a 9year period (1940-1948), after which followed deep depressions.

The

fluctuation

was not 2-4-fold but

16-fold. Increases in

numbers

1051
repeated after 2-3 years (1940, 1943, 1945, 1948), and in a few
cases, in the year, after an increase in

numbers of rodents (P.B.

Yurgenson).
Field characters. In nature, the solongoi differs from ermine
(in

summer)

in denser, fluffier tail in its

without a black

tip.

It

differs

from the kolonok

smaller dimensions and paler color (P.Yu.).

Practical Significance

The significance of solongoi in fur trade is not great. In Kazakhstan, its catch constitutes 0.2% of the tanned fur. Nowhere is it hunted
commercially.
It is

captured incidentally, but not specifically

is

trapped. In the Balkhash region, the solongoi

harmful to coma harm-

mercial hunting, destroying muskrat and pheasant, and damaging

the skins of muskrats in traps. Here,
ful predator.
it

may be considered

and for ecosystem health, it may be considered it destroys harmful rodents. However, this benefit is not great, since solongoi are encountered predominantly in uninhabited or weakly populated areas. For the same reason, the harm it causes by attacking domestic poultry is insignificant. In pre-war years in the USSR, about 50,000 animals were captured
In agriculture

a useful animal since

annually (Stroganov, 1962) (P.Yu.).

KOLONOK

[SIBERIAN WEASEL]

Mustela (Mustela) sibirica Pallas, 1773

1773. Mustela sibirica. Pallas. Reise
Reiches,
Altai.
2, p. d.

versch. Prov. d. Russ.

701. Tigeretsk Post near Ust-Kamenogorsk,

1844. Mustela

itatsi.

Temminck. Fauna Japonica, Mamm.,

p. 34,

t.

VII, Fig. 2. Japan.

1871. Putorius fontanieri. Milne-Edwards. Rech.

pi.

Mamm.,

p.

205,

61, Fig.

1.

Pekin [Beijing], China.

1904. Mustela sibirica miles. Barret-Hamilton. Ann.

Hist. 13, p. 391. Dauriya, Trans-Baikaliya.

Mag. Nat.

\9\\. Mustela manchurica. Brass.

490. Manchuria.

Aus dem

reiche der Pelze, p.

1052
701

1911. Kolonokus sibiricus australis. Satunin. Izv. Kavkazskoyo

muyzeya,

5,

p.

23. Zhiryakovo,

Antropovsk. vol[ost].,

Tyumensk

county.

1926. Kolonocus sibiricus coreanus. Domaniewski. Ann. Zool.

Mus. Polon., 5, p. 55. Seoul, Korea. 1934. Mustela {Kolonocus) sibirica charbinensis. Lowkashkin
[Loukashkin]. China.
J.

Sci.

and Arts, 20,

p.

49. Krestov

Island on Sungari river near Harbin, Manchuria (V.H.).

Diagnosis
General color monotone and evenly bright, ocherous-reddish or

rusty-brown tones; venter same color as dorsum or slightly lighter;

on sides dorsal color gradually merges into ventral coloration without noticeable boundary. End of muzzle and region near eyes coffee-brown ("mask"), and on lips and chin, a sharply outlined white area. Legs no darker than trunk. Postorbital constriction is elongated, its lateral outlines almost parallel, no distinct isthmus directly behind supraorbital processes. Dimensions condylobasal length of male skull more than 55 relatively large mm, of female, more than 49 mm (V.H.).

Description

The kolonok has

a long stretched-out

body on

relatively short legs;

with, however, a general appearance of being heavier than solongoi,

and especially ermine and weasel. In this respect, it seems to occupy an intermediate position between these forms and the larger members of the genus mink and especially polecats. This is also associated with the relatively large measurements of the kolonok. The head of the kolonok is relatively small, narrow and elongated. Ears are broad at base, but not tall, and in winter, barely protrude

from fur. Length of tail with terminal hairs constitutes about half of body length. Winter fur very dense, soft and fluffy. Length of guard hairs reaches 30-40 mm; underfur dense and close-fitting. General color
of entire animal monotone, bright reddish-ocherous or straw-red,

sometimes, orange or peach tone noticeable on skin. This color

especially bright on back, on sides a bit paler,

somewhat

paler yet

on abdominal side. Neck and throat colored like abdomen, outer side of claws same color as dorsum of body.

1053

Fig.

250. Kolonok, Mustela (Mustela) sibirica Pall. Sketch by A.N. Komarov.

On

anterior part of head, covering the circumference of the

eyes and forwards to the end of the nose, a dark coffee-brown

sharply outlined spot

"mask".

Upper and lower

lips

sometimes

have light ocherous highlights. Their color sharply delimited both from dark color of mask and from adjacent ocherous-red areas. In some individuals, white color covers bare tip of nose in form of narrow band extending from sides and back [of head] or from
sides only. Sometimes,
throat.
702
It is

one or several white small spots occur on

Tail usually brighter than

back

bright-red above and below.

is

covered by long, dense, loose-fitting hairs and

all

very fluffy,

fluffier than all

other species of genus, including polecats and

in

minks. Tail thickest of

towards
tapered.
its

middle

part, gradually thinning

at tip,

base and rapidly coming to naught

part of

which

is

Lower

paws covered by dense,

elastic

silvery-

straw hairs, through which, however, callous digital pads located

beneath the hairs show. Dorsal paw has same color as back.
In

summer
it

fur,

whole trunk covered by

shorter,

coarse,

quite close-fitting hairs.

ting; therefore

On

tail,

hairs relatively short, close-fit-

appears thin, of equal thickness along entire length.

1054

Color almost completely monotone over whole body. General tone

of back dark with intense ocherous-brown, and with middle of back only somewhat noticeably darker. Sides same color as back, ventral surface of body only slightly lighter than dorsal. Color of throat only slightly more intensive than abdomen. Upper side of fore legs often slightly darker than hind, which have color of sides. Tail, dorsally and ventrally, has same color as back. Lips (upper and lower), chin, and often narrow stripe around

bare part of nose are a pure white or dirty-white color, or slightly

ocherous, but always sharply demarcated from color of surrounding parts of body. Anterior part of muzzle darker than remaining
part of head, but this "mask", in contrast to that in winter,
is

not

so sharply limited;

its

posterior border

is

completely unclear and

color of "mask" here gradually transitions to color of top of head.

Individual variation in color has small range, manifested in

general tone, size and intensiveness of

mask

color and white field

around mouth. Sexual differences

mals are clothed in downy brown

in color are absent.

fur,

Young

ani-

darker than in adults. First

winter coat does not differ from fur of adults. Geographic

variation in color of kolonok within boundaries of

USSR

almost

unexpressed or expressed very weakly and revealed only in some

deviations in

mean

intensity of general color tone.

Among

kolonoks albinos are encountered

complete or

partial
tail,

(Zalesskii, 1970), white with light red guard hairs

on back and

and several other mutational variants. "Giant kolonok" mentioned

below have color which is not normal. The kolonok skull occupies, in several characters, an intermediate position between the skulls of ermine and mink. It is larger and more elongated than ermine and is somewhat flattened compared with the mink skull. In general outlines, the skull is narrow and elongated, considerably narrower and longer than solongoi, and the brain case is elongated and weakly swollen. Mastoidal width usually less than half condylobasal length of skull and significantly less than distance from anterior edge of foramen magnum to anterior point of interpterygoid notch. Region of postorbital constriction elongated, its isthmus almost

unexpressed or
of brain-

weakly expressed, and outlines of

can be noted, then
Its
it is

lateral sides of this part

case almost or more or less parallel. If something of an isthmus

not situated close to supraorbital processes.

narrowest part

is

located

away from

the line uniting supraorbital

1055
processes, not less than half the distance between the outer ends of

these processes.

703

Fig. 251. Skull of the kolonok,

Mustela (Mustela) sibirica

Pall.

1056
Infraorbital
to
it.

foramen

less than alveolus of

upper canine or equal

Interpterygoidal area does not have form of a forwardly

directed angle, and does not evenly narrow anteriorly. Auditory

bullae elongated, their width constituting about half their length or

704 less. Distance

somewhat
their

between inner borders of anterior parts of bullae Distance between bullae in anterior part less than width of hard palate at anterior edge
less than in posterior part.

of interpterygoidal notch.

Zygomatic width approximately equal to skull width in masZygomatic arches elongated and weak. Protuberances, crests, etc. of skull relatively weakly developed,
toid region or slightly more.

but generally stronger than in solongoi. Sagittal crest clearly

noticeable in posterior part of cranium and in anterior,
defined; occipital crest well developed.
is

weakly

Upper

profile of skull

uneven; an elevation is noticeable on it in posterior part of cranium and prominence in frontal area (interorbital region) is obviously convex.
facial region

From

the frontal area, the line of the profile of the

descends steeply downwards.

Teeth, especially canines, relatively large and strong.

On

the whole, the kolonok skull represents the skull of a spe-

cialized predator and does not itself bear the infantile characters

known in the genus which characterize the solongoi or at least of some of its races. This also applies to weasel and ermine to a lesser degree. In some respects, the kolonok skull is even more
specialized than the

mink

skull.

Age and
for other

sex differences of the kolonok skull have not been

especially studied. Generally, they correspond with those mentioned

members of the genus Mustela (see description of ermine). The female skull is considerably smaller than the male, somewhat lighter and with less defined protuberances, crests, etc. In young the cranium in the interzygomatic region, in particular, is more swollen. Individual variation of the skull does not exceed
"normal" limits for other close species, except the weasel. Geographic variation of the skull, within the boundaries of our country, is

not at

all

or

is

very weakly emphasized, and bears only a

typical of the majority of

quantitative character (general measurements).

The

structure of the os penis

is is

species in the genus

the

distal

end

bent upward in the form of

a hook. Also characteristic are several bends in the basal shaft of

the penis, in the distal part; thanks to

which

all

of

it

is

directed

1057

somewhat upward. The quite large dimensions of the bone are more than in the species described above, comcharacteristic

pared to the general dimensions of the animal.

In dimensions the kolonok is considerably larger than ermine and solongoi and approaches ferret and mink, but is somewhat inferior to them. Body length of males (47) is 280-390 mm, of females (38), 250-305 mm; tail length of males is 155-210 mm, of females, 133-164 mm; length of hind foot of males is 53.6-68.0

mm,

of females, 48-52

mm;

ear length of males

is

22.0-29.5

mm,

of females, 19.0-23.0 mm.^^

Condylobasal length of skull of males is 58.0-67.3 mm, of mm; zygomatic width of male is 28.7-35.7 mm, of females, 26.4-32.1 mm; interorbital width of males is 11.7females, 49.8-62.7
13.9

mm,

of females, 10.5-13.0

mm;

mastoid width of males

is

26.8-31.0

mm,

of females, 23.0-27.0

mm.

Os penis length is 32.0-35.8 mm. Weight of male is 650-820 gm, of female, 360-430 gm male (probably juvenile) is 395(Stroganov, 1962), and also less
485 gm, female about 300 gm (winter materials of Z[oological] M[useum of] M[oscow] U[niversity]).
In western Siberia (Barabin steppe), sometimes, but very rarely,

giant kolonok occur with significantly larger dimensions than usual.

If the length

of a commercial skin of normal kolonok from the tip

tail

of the nose to the end of the

with
it

its

terminal hairs reaches

only 59-60 cm, then in giant animals,

ranges between 80 and 88

in "giant"

cm. As well, color varies greatly

705

undercoat

kolonok

is

almost white (dirty yellowish-white or light-grayish), and paws and end of tail are dark-brown (Zalesskii, 1930; Zverev, 1931). Therefore, several characters of the steppe (white) ferret are well ex-

pressed in them. Apparently, this

the kolonok, but a hybrid

is

not an "exotypic" mutation of

ferret (M.

between kolonok and steppe

eversmanni). These individuals are obtained in the region where

both species live not only in one territory but also in one and the same biotope. Their large measurements are, apparently, an illustration of heterosis, "Giant

kolonok" are known only from skins

310^30 mm,
tail

^'From other data (Zverev, 1931; western Siberia), body length of males (55) is often 350-370 mm, of females (60), 300-360 mm, often 260-310 mm;
is

length of males (50)

often

140-220 mm, often 160-210 mm, females

(55),

130-210

mm,

150-170 mm.


1058
prepared as furs, and their skulls especially were not studied in
detail.

"Giant ferrets" are

known

in detail (see

below) (V.H.).

Systematic Position

As already mentioned above

in

(see section "Systematic Position"

solongoi chapter), an adequate bases for affiliating kolonok

and solongoi is absent, the more so as they do not happen to be considered the_ closest species in the genus, combined in one group as opposed
to others, etc.

Other qualities than

in solongoi

color type (no contrast in color of upper and lower body either in

winter or in summer) and

its

geographic and seasonal variability

are the essential indices of the deep differences

species.

between these

The closeness of features of kolonok

In purely morphological features, the

to

mink

are without doubt.

kolonok obviously differs however, information concernfrom ferrets (subgenus Putorius); ing hybrids between these species occurring in nature (see above) point to a close connection between these seemingly separate species. This is, once more, confirmation of the homogenity of the genus Mustela, and at the same time an example of the complicated inter-relationships of
all
its

species.

It is

difficult to array

them

in

successive order, and quickly becomes necessay to think

about a quite complicated arrangement around a radial [pattern].

The order of species which is closer here as the most natural (V.H.).

to the traditional

one

is

adopted

Geograpliic Distribution

and Central Asia, the southern and middle zone of Siberia and the Far East, and in the eastern European part
In Southeastern, Eastern

of the

USSR.
in the Soviet

Geographic Range

Union.

Vast, constituting the greater part of the northern half of the range,

and occupies the southern and middle zone of Siberia and the Far East as well as the eastern European part of the USSR. In the European part of the USSR, the range of the kolonok forms a projection having as its base the Ural mountains, and
directed westwards. In the Urals, the northern border passes just

1059

below 63 N.

lat.

from whence

it

goes to the southwest through the

southeastern and southern outskirts of

of the tributary of lower Sukhona

east,

the Luza (Oparino

Komi

district to the

source

to the north-

passes below 6020' N.

dle Vetluga.

and Letka and Ob'yachevo to the north of Kirov), where it lat. (Ob'yachevo), and thence to the mid-

From

here, the border line apparently passes generally

is

along the Vetluga and Volga to Kazan, and then

directed to the

southeast, covering a great part of Tatariya, the northern- and

707

northeasternmost outskirts of Kuibyshev district and Bashkiriya, and passes onto the southern Urals^. In the extreme south of the Urals, the kolonok is distributed over the whole montane region and is also encountered in the forest-steppe between the Sakmara and Ural (in the area between Kuvandyk and Saryktash') and even in the Guberlinsk melkosoposhnik [small hills] (the inner part of the angle formed by the winding Ural) to Guberlinskaya (somewhat westward of Orsk) and in the floodland of the middle course of the Ural.
In Siberia, the northern border of
lat. in

kolonok range from

**

N.

the Urals, crosses the Ob' river, passes to the upper Pur and

then suddenly ascends to the north on the lower Taz, almost reaching the Arctic Circle here or, perhaps, even a
the east, the border
still

little

beyond

it.

To

ascends to the north, passes to the upper

Turkhan and the upper Kheta, crosses the Yenisei between Dudinka and Potapov (above Dudinka) and passes east of Yenisei in the
region of Khantaiskoe and Ketoi lakes (69 N.
lat.).

This

is

the

northernmost confirmed point inhabited by kolonok.

Farther, the border, in an unclear way, suddenly descends south-

ward and

on to the border of Yakutiya approximately at 63 N. lat., and at that latitude, crosses the upper Vilyui in the region of the mouth of the Chona (Maak, 1859; D. Ivanov). Still more to the east, the border passes to the
then, returning to the east, goes

middle course of the

N.
lat.)

left tributary

of the Vilyui

the Markha (64

and reaches the Lena

to the north

of the mouth of the

Tatariya, the kolonok

'"According to V.A. Popov (Kazan), during the whole time of investigations in was never found west of Chistopol'. This same author also

considers that the animals which were caught in the Vetluga basin (Dryablovo,
tributary

Krasnye Baki; Formozov, 1935) were themselves not naturally occurring animals, but were kolonok introduced into the Vetluga basin. At the same time, data on catch of these animals (1927-1932) and the known data on introductions of kolonok into the Trans-Volga (Semenov city on the Kerzhenets 1937; Puzanov, Kozlov and Kiparisov, 1955) do not offer a basis for this assumption. From the 40' s, kolonok were not observed in Gorki district.
of the

Vetluga

the

left

Kaksha,

>a

.^

1061
Vilyui at 65 N.
right

lat.

Thence,

it

descends

to the south

along the

bank of the Lena

right tributaries

valley, covering thereby the lower courses

of

its

and thus goes

to the nnouth of the

its

Aldan.
exthe
at

Along

the Aldan, also covering the lower courses of

right tribuit,

taries, the

border goes to the Allakh-Yun' River and along

tends beyond the border of Yakutiya (D. Ivanov).

From

Allakh-Yun', the boundary passes to the mouth of the Okhota the Sea of Okhotsk (about 5910' N. lat.; V.P. Sysoev).
individuals,

Individual cases of captured kolonok, apparently, dispersing

were recorded

in

of the above-mentioned line

east of the

along the Tyung

Yakutiya and somewhat northward in its upper course,

along the Lena in the direction of Zhigansk and somewhat north-

mouth of

the Aldan.

all

South of the specified line, the kolonok is distributed through of Siberia, and in the expanse extending from the Pacific Ocean

to the southern Altai its range passes

beyond

the southern frontier

of the

state. It is

absent only in the south of southeastern Trans-

Baikaliya, and particularly south from the line going from the

Argun' a

little

south of Nerchinsk Zavod, across the Kalga to

left bank of the Onon, and thence southwest to the state frontier. In southern Altai, the Lake, but is kolonok extends south to the latitude of

Borzya, to Olovyannaya, and then to the

absent in the Zaisan depression.

To
north

the west of the Altai, the kolonok

is

met with

in the

Kalbinsk
south
of

Altai (left

bank of the

Irtysh), but farther south the

border goes

again,

generally

along
is

the

Irtysh,

although

Semipalatinsk, the kolonok

encountered locally on

its left

bank

(Zharminsk region, for example). Farther, in northern Kazakhstan, the animal exists in the northern part of Pavlodarsk district (Irtysh region below Pavlodar) and in Severo-Kazakhstansk district, it is met with among the insular forests of Kokchetav and even the
former Akmolinsk
a
little to

district.

South of here,

it

penetrates to the area

little

the north (Alekseev forestry) and even a

southeast

of Tselinograd (Vishnevsk region). Farther, the range boundary

includes the northern and western parts of Kustanai district (east

of Tobol), and proceeds to the Ural [river] in the Orsk region.

On

the Shantar and Kuril islands, the kolonok

is

absent.

On
that

Sakhalin, fossils of Siberian weasels were also absent, but in 1932,

the Japanese weasel

was introduced

to this region,

and from

time, a quite permanent population has, apparently, been founded

in Sakhalin, at least in the south.


1062

The range of

the kolonok in the last century has undergone

in a series of places these

considerable changes and

continuing at present. In several places in central Siberia,

changes are Yakutiya

and the Far East, the kolonok

westward. Thus, even
708

is

extending
is

its

range, though not

very intensively. Particularly notable

in the 70' s

the kolonok' s colonization

last

and 80' s of the

i.e.

century

it

was absent

in the Urals.

The kolonok occupied

range,

the

whole Euro-

pean (Cis-Ural) part of

it

its

a very significant territory,

only in the present century. In the northeastern part of Tatar

ASSR,
it

appeared around 1910.

On

the Vyatka, in Kirov region,

it

was

first

noted in 1926; in 1927

penetrated to the eastern parts of

it

Gor'ki district (Vetluga basin);

sources of the Luza

was

rarely encountered on the

right tributary of the lower

Sukhona, and by

became "almost common" here (Ob'yachevo, Letka). In some places, in a wide belt along the periphery of the range, sharp fluctuations in the boundary and numbers were noted in
1949
it

kolonok. This particularly applies, apparently, to the westernmost part of the range, where the distribution of Siberian weasel is
sporadic, the greater part of the boundary
is

not only very badly

understood, but

is

evidently also in general

little

defined and sub-

jected to strong fluctuations. Thus, for example, in

lying within the border,
it

Perm

district,

found everywhere, except in the northwesternmost part. Up to 1925 the animal was very rare in the Tatar Republic. Then in 1925-1928 it became a mass [abundant]
is

species and noticeably spread to the west, but in the 40' s

it

again

became

rare.

In Gor'ki district, appearing in

1927, the kolonok

declined in the course of several years, and again disappeared, and

since the 40' s has not been recorded.
In Novosibirsk district, the kolonok

up

to 1940, but in 1955, the southwestern

was found in all regions boundary of its range

extended from Ust'-Tara on the. southeast to the northern part of Lake Ghana, and thence, arching towards the north to Kochka region which lies southwest of Novosibirsk. In northern Kazakhstan,
it

becomes more common, inhabiting

appears rarely, and transgressions do not occur every year, then the territory and being caught

regularly, and again completely disappears for years.

These

fluc-

tuations in the range boundaries of the kolonok in Kazakhstan

1063

in

Kustanai

district

and Tselinograd

territory are, apparently, very

sharply expressed.^'

The boundary and numbers,

apparently, change noticeably along

the Irtysh. Therefore, the outlined range also areas of transgression and greatest

is maximal, and includes movements. Colonization was particularly intensive, first of all, in the European part of the USSR in the 20' s, and perhaps the beginning of the 30' s. Later, it weakened, and ceased; the boundaries of the range more or less stabilized, and in some places, it again decreased. Apparently, the situation was also nearly the same in southern Siberia and in Kazakhstan. It is possible that in western Siberia, there were also local extensions of the range boundary to

the north; however, data about this are absent.

It is

possible, apparently, to think that the previous coloniza-

tion of the kolonok,

which was observed

in the past decades, as a

earlier.

continuation of a process begun

much

This belief lacks

supporting data, but

Siberia, or at least
is
its

it

is

probable that the kolonok colonized

parts, in the past century. This
its

more northern

indirectly

shown by

the characteristics of

range, which poorly

fits

into the

usual landscape-geographic framework, and by

several other circumstances, in particular weakly expressed geo-

graphic variation. However, there

is

considering that the kolonok

Siberia in

was generally absent the 18th century and that it was only found

no basis (Laptev, 1955)^^ for in western

east of the

Yenisei (Bobrinskii, 1946).

"This phenomenon
Kazakhstan.
is

also well expressed in several other species, the southern

distribution boundaries of

which extend south

to

western Siberia and northern

which is 1944 and others). This particularly applies to the boundaries of the range in Siberia. In particular, it is sometimes shown that the kolonok is distributed "over all of Yakutiya, " its occurrence near Verkhoyansk is also mentioned. In composing notes on the range, the following works were used: Maak, 1859; Shrenk, 1859; Sabaneev, 1874; Kashchenko, 1898; Ognev, 1931; Tugarinov, Smirnov

"The described range of

the kolonok differs significantly from that

usually described and illustrated (Bobrinskii,

and Ivanov, 1934; Adlerberg, 1935; Formozov, 1935; Heptner, 1936; Podarevskii, 1936; Lavrov, 1946; Kuznetsov, 1948, 1952; Popov and Lukin, 1949; Shvarts, Pavlinin and Danilov, 1951; Kirikov, 1952; Sludskii, 1953; Kiparisov, 1955; I. Laptev, 1958; Parovshchikov, 1959; Afanas'ev, 1960; Stroganov, 1962; Puzanov, V. Kozlov and a series of other authors. Chief among those providing original material can be selected V.N. Skalon (West and Central Siberia), P. Zimin (Chitinsk district), Bel'sk (Primor'e Territory), A. Samsonov (Amur district), S. Anashknin (Buryat ASSR),
D. Ivanov (Yakutiya) and V.P. Sysoev (Khabarovsk Territory).
In

particular,

through these materials, the whole boundary in Siberia was revised. V.A. Popov (Tatariya) and S.P. Chashchin (Perm district) provided their data on the Euro-

pean part of the USSR.

1064

Kolonok were introduced

1937
in the

for the sake of acclimatization in

Semenov region

of Gor'ki district (individuals from

West

Siberia) and in 1941, was introduced near Przheval'sk (Dzhetyoguz region, Chu-Bulak ur.,* individuals from Krasnoyarsk

Territory).
709

Geographic Range outside the Soviet Union

This
is

vast, with

complicated outlines, but

still

poorly studied.
it

The kolonok

exists in the

Mongolian Republic, where

occupies

Fig.

253.

Natural species range of kolonok, Mustela (Mustela) sibirica Pall.

V.G. Heptner.
*Contraction for urocliishche, meaning "small village"

Sci. Ed.

1065
Pri-Kosogol', Khangai and Kentei; on Korean Peninsula and in China northeastern (former Manchuria), and to the south including Guandong and Yunnan, in Tibet, in the Himalayas, to the west including Kashmir, and in Nepal and upper Burma. It exists on the Japanese Islands*, Taiwan and Java**. There is no information about the occurrence of the kolonok in Indochina, and therefore the range of the species shows a strong and complete disjunction south China and upper Burma on one side and Java on the other. That is how it is usually described (Pocock, 1941; Ellerman and Morrison-Scott, 1951). It is possible that this questionable gap is filled by other forms, which are considered separate species, in part, possibly, M. nudipes, which is close to M. sibirica and which inhabit Malacca, Sumatra and Kalimantan (Borneo). The elucidation of the actual picture demands further investigation, not only on the faunistics of Southeast Asia, but also the systematics of the kolonok (V.H.).
the

northern montane region

710

Geographic Variation
forms have been described for kolonok, among them some from our territory or neighboring territories. At the same time, as shown by the large collection materials Z[oological] M[useum of] M[oscow] U[niversity], geographic variation of the species within the borders of the USSR, i.e. the vast area extendseries of

including

ing from the Volga Basin to the Pacific Ocean,

is

very small

small
is

number of

real

forms and amplitude differences

its

in characters

insignificant. This leads us to think that variation in the species

is

generally insignificant, and

presentation

is

exaggerated

(Ellerman and Morrison-Scott, 1951).

The scheme
1962)
is

recently suggested for our territory (Stroganov,

is

mainly based in part on exceptionally subtle characters, the reality of which is highly doubtextremely fragmented and
ful.

The recognition of
in

the concept natio (for the

form

australis),
it

which

essence

is

lacking in real content, does not

make

more
itatsi

convincing. In exactly the same way, separation of the form

into an independent species (see

below)

is

also not well founded.***

Geographic variation

in

our kolonok

is

considered very insignifi-

*Considered by some to be a distinct species, M. itatsi Sci. Ed. **Considered by most to be a distinct species, M. lutreolina Sci. Ed. ***Contradictory? Sci. Ed.

1066
cant even by furriers (Kuznetsov, 1941). Several East Asiatic insular forms, in particular itatsi are better distinguished* in natural

conditions of occurrence outside our country.

Within the borders of our country, two forms

guished.
1.

may be

distin-

Siberian kolonok, M. (M.)

s.

sibirica Pallas,

1773 (syn.

aus trails, miles).

Measurements comparatively small. Color of winter fur not vey bright, yellowish-red.
Condylobasal length of male skull (33) is 58.0-M61.7-63.5 of females (15), 49. 8-M52. 8-56.3 mm; zygomatic width of males is 28.7-M32.2-35.2 mm, of females, 26.4-M27. 8-29.6 mm; interorbital width of males is 11.7-M12.5-13.2 mm, of females,

mm,

10.5-Ml 1.0-12.2 mm; mastoid width of males is 26.8-M27.5mm, of females, 23.0-M24.3-26.1 mm." Found in eastern European part of the USSR and in all Siberia eastward to Zeya basin.
28.7

Outside the

USSR found
its

in the part

of the Mongolian Re-

in extreme western China (former Manchuria). In some parts of the range, there can be noted a tendency towards deviation in the character (quality) of the fur coat

public contiguous to

range and, possibly,

parts of northeastern

(density, softness) and in its color (West Siberian lowlands, Yakutiya, Bashkiriya). These deviations are more noticeable in

Trans-Baikaliya, where kolonok are somewhat brighter and, apparently,

themselves represent a transition to the next form. All these

deviations, even those in the Trans-Baikal population are,

ever,

how-

weak and

their territorial limits are not clear

and do not offer

(in

a basis for separation of independent geographic forms.

2.

Far Eastern kolonok, M. (M.)

s.

manchurica Brass, 1911

in

our literature, the

color lighter, red.
711

name coreana is accepted). Average measurements somewhat larger than

nominal form;

Condylobasal length of male skull (16) is 63. 8-M65. 4-67.3 of females, 57.2-M61.2-62.7 mm; zygomatic width of males is 32.3-M34.3-35.7, of females, 27.0-M30.5-32.1 mm;

mm,

inter-orbital

width of males

is

12.3-M13.2-13.9

mm,

of females,

Contradictory?
ments given above

Sci. Ed.
this

"Measurement of
(p.

and the next forms from Stroganov (1962). Body measure-

1057), from Zverev (1931), belong to this form.

1067

11.0-M12.7-13.0
14.0

mm;

postorbital width of males

is

13.2-M13.6-

mm,
is

of females, 12.0-M12. 7-13.3

mm;

mastoidal width of

males

This form
Primor'e.

28.2-M29.5-31.0 mm; of females, 24.0-M25. 0-27.0 mm. is found in Priamur'e to the west to Zeya [river],

Outside the
ria),

USSR in

northeastern China (former

Manchu-

precise limits

unknown.

well defined form. Apparently, identical to the Manchurian

[form] and must carry the

name recognized

here.

Comparison of
however, been

our Far East kolonok with Chinese and Korean, the names of which

might be applied

to ours (see

synonymy) has

not,

made and
plied.
3.

name manchurica, therefore, conditionally apIt is probable that the name fontanieri may be more proper. Japanese, or Sakhalin, kolonok^'*, M. (M.) s. itatsi Temminck,
thus, the

1844.

Measurements somewhat smaller than in Far Eastern kolonok, M. (M.) s. manchurica. Color of ventral surface of body somewhat
lighter than dorsal, limbs slightly darker than trunk.

Skull form as in other races, but frontal area

bulging, inner border of auditory bullae

posteriorly,
rior part is notably

somewhat more somewhat separated

in their poste-

and distance between auditory capsules

more than anterior. Part of hard palate lying behind the [last] molar somewhat narrower than in other races. Body length of males (9) is 510-550 mm, of females (2), 341-465 mm; tail length of males is 145-165 mm, of females, 93101 mm; length of hind foot of males is 57-65 mm, of females, 39-46 mm.
Condylobasal length of male skull
males, 47.6-57.9
is

57.2-61.9

mm,

of fe-

zygomatic width of males is 31.3-33.2 mm, of females, 28.9-32.0 mm; interorbital width of males is 12.012.8 mm, of females, 9.1-12.0 (after Stroganov, 1962; supple-

mm;

mm

mented by material of Z[oological] M[useum

U[niversity]).

of]

M[oscow]
in 1932.

Found

in Sakhalin, only in south.

Acclimatized there

Outside the

USSR in
Iki).

the Japanese Islands (Hokkaido, Honshu,

Shikoku, Kyushu,
'^It

was suggested (Pereleshin, 1957; Stroganov, 1959, 1962) that this form be name "itatsi" for us. This name must be decidedly rejected. There is no reason at all to litter the Russian language with a word completely alien to it,
given the
especially a non-declinabie one.

1068

well-distinguished form, differing

latter

more strongly from named

forms than the

do from each
to
it

other.

Note. The relatively sharp differences between Japanese kolonok

and Siberia turned attention

borders of the

USSR

(Pereleshin, 1957).

when first recorded within the Somewhat later, Stroganov

(1960, 1962) decisively affirmed that the form itatsi is itself an independent species differing from M. sibirica. According to the

words of
(12 in

this

author and the majority of characters noted by him

particular, 5 out of 9 of

all), in

"sharp hiatus" exists between the form

which are craniological, a itatsi and those of the

of]

mainland.

Materials

in

the

Z[oological]

M[useum

M[oscow]

U[niversity] from Sakhalin (collected by A.I. Gizenko), however,

show

that the features distinguishing the

are far

from such

as those described

form itatsi from Siberian and illustrated by Stroganov

is

(1960, 1962).

clear hiatus

between

absent in any one. All are

either not developed or are extremely variable

and broadly over-

lapping. In particular, differences mentioned as of fundamental

important and striking

position of the anterior edge of the audiits

tory bulla in relation to the articulated groove (anterior edge of

bulla found a distance half the bullar length from the articulated

groove

in itatsi

and

at

a distance of one-fourth

length in sibirica)

as well as dimensions
712 at all in material of
ters].

and form of the bulla

are not demonstrated

the diagnosis

ZMMU.

The same
is

applies to other [charac-

Some

bulging of the frontal area

bulla,
as

noticeable (weak) and a

well expressed difference in the position of the inner outline of the

boney auditory

mentioned
itatsi

in

above.
is

Therefore, to believe that the form

inhabiting Sakhalin
It is

an

independent species
its

is

without foundation.

a true kolonok and

somewhat

greater differences are, apparently, explained

insular origin.

by its The conclusions of Stroganov (1960, 1962) were

based, apparently, on accidentally selected, atypical materials.*

Data on color dimensions could not be verified and are after Stroganov (1962). They require re-examination. The structural
peculiarity of the posterior region of the hard palate

was not men-

tioned earlier.

Recent work now

supports Stroganov's opinion

Sci. Ed.

1069
Outside the borders of our country, the following forms are
usually mentioned:
1)

M. (M.)
to

s.

subhemachalana Hodgson, 1837

Bhutan; 2) M. (M.) s. canigula Hodgson, 1942 Tibet; 3) M. (M.) s. hodgsoni Gray, 1843 Kashmir and western Himalayas

Nepal

Himalayas from

from Kam to Garwal (Garhwal); 4) M. (M.) s. davidiana MilneEdwards, 1871 Southeast China north to Hubei, Taiwan; Northern China, 5) M. (M.) s. fontanieri Milne-Edwards, 1871 Shandong, Chihli, Shansi and Shensi; 6) M. (M.) s. moupinensis Milne-Edwards, 1874 Sichuan, Gansu and Yunnan in China; Burma; 7) M. (M.) s. manchurica Brass, 1911 Manchuria; 8) M. Korean Peninsula; 9) M. (M.) (M.) s. coreana Domaniewski, 1926 charbinensis Loukashkin, 1934 Manchuria; 10) M. (M.) s. s. quelpartis Thomas, 1908 Quelpart Island in East China Sea; 11) 1924 Tanageshima and Yakushima isM. (M.) s. sho Kuroda, lands (southern Japanese archipelago) [= Ryukyu]; 12) M. (M.) s. Oshima Island, Japan. asaii Kuroda, 1943

is

The

list

given

obviously inflated, probably approximately

in

twice. For

some forms, deviations

which are

characters of the facial

pattern, the majority of

fairly variable, small differences in

color intensity etc. are considered to be diagnostic characters (V.H.).

Biology
Population. The range of the kolonok
is

very extensive, but

it

exists in entirely different densities in relation to the great diversity

of landscapes and, consequently, conditions of habitation.

Figures of [fur] preparation do not reflect these particularities. The

southern regions of Siberia and the Far East are richest in kolonok.

The
the

latter

produces more than 1/3 of the skins prepared

In the Gorno-Altai
(in

in

USSR.

frequency of tracks found

10

Autonomous District, the highest km) in a series of years ranged

from 2.85
a

to 6.6,

and the lowest

from 0.01

to 0.22. In the taiga

of the Sikhote-Alin' range, the number of tracks encountered along

10-km route ranged from 24.1

it

to 38.8 in half the cases, but

sometimes

decreased to 2.1 (Yu.A. Salmin and V.D. Shamykin).

stability of

Population and distribution of kolonok are determined by food

abundance (mouse-like rodents and water voles), the

taiga

stocks, interrelationships with competitors (in dark coniferous

with

sable; in taiga river valleys

with

ermine and also

1070

American mink;
ermine;
at the

at

the upper edges of forests

steppe border

with

solongoi
it

and

steppe polecats) and also the

willingly

characteristics of

snow cover. Habitat. Kolonok habitats

are varied. In the taiga,

holds to valleys of forest rivers and creeks with dense thickets as well as the rocks and rock slides overgrown with bushes, forest

marshes (sogra) and,

dense growth of saplings. kolonok prefers dark coniferous taiga with larch stands. However, these are secondary habitats whence it penetrates after the extermination of sable, and from
after old burns,

In the Altai mountains, the

crowd them out. In spruce-fir forests of the Sikhote-Alin' range, the kolonok is most numerous in mixed nut pine broad-leaved taiga, broad-leaved forests of the Manchurian
which
after a time sable again

Fig. 254.

Mountain creek

typical

habitat of kolonok.

"Kedrovaya Pad'

" preserve,

southern Primor'e. October 1963. Photograph A.G. Pankrat'ev.

1071
713 type

to the

and in coastal oak forests. It is met with from the sea coast upper limit of the forest, but in the latter, it is rare. In the forest-steppe of Siberia and the Far East, the kolonok

inhabits birch and aspen clearings, reed thickets

"flood-lands"
steppes
It

along the banks of steppe lakes and rivers, and

with shrubby growth, and rarely

meadow

pine forest islands.

penetrates

the depths of the taiga immediately after agricultural developments

and human settlements, which is explained by concentrations of mouse-like rodents in these places. It often inhabits villages and
even
cities

where on the one hand

it

destroys mice and rats and on

It

the other causes

damage

to

domestic fowl.

avoids open places.

Food. According to type of food, the kolonok belongs to a group transitional between the typical mouse-predators (weasel) to
the polyphagous predators (true martens). Cold-blooded vertebrates,

invertebrates (insects and molluscs) and plant food are not a

consistent element in
its

ration

and are not encountered every-

where. Rodents of small and moderate size constitute the basis of

714

Table 67. Geographic variation in foods of kolonok

(%
a

of occurrence)

>
(Fetis

JS.

>

i! 5S ^

^
Type of food
55 C/3

i 00 <u
.

en

2
Al
931)

stem

2)
194

w
Zverev

^
62)

N^^

TO

ppe,

2
ern

1956

a^
S

ppe,we;

kaliya

>
1)

-^
=

-<

.2

'^

<
-

H
a

(U

?
(U I[1,

tS =

H
Water vole [Arvicola
terrestris]

CQ ^ ra

-o
rt

X) . CO t^

> 2 u E

1
u ^ .Q 00

'" fl
OX)

2
2
>
ce

7,

a 1
^^
u.
11-

-5

g
_4
v^"

'

= t;
Z
CL

^ XI 2 z.
r^

-J

d-

H -

^ ( ,t'='

M
.2?
'

-s

iS ro 0\

"

/5

36.6
15.0

56.1

46.0
15.0

1.8

7.0

68.7

Small voles and mice Altai zokor

[M. myospalax]

Great jerboa
[Allactaga major]
Siberian

chipmunk

[Tamias sibiricus]
Pikas
Insectivores

130

15.6

6.6
3.3

0.8

43

50.1

20.0 21.0
14.0

24.6

0.9
2.7

2.0

9.6

38.2
3.5

33.8

Birds
Fish

4.6
1.7

15.0

13.3

15.0

Carrion

108

6.4

52

29.0

13.6

9.4 4.4
10.1

406

0.5

Amount of

data

60

684

examined

1072

kolonok food everywhere. Species composition and ratio of each species depend upon the local conditions and competitive relationships with other carnivores. Geographic variation in primary

714 Fig. 255. Bank of "Kedrovaya Pad'"

montane creek
preserve,

characteristic

winter habitat of kolonok.

southern Primor'e. A.G. Pankraev.

March

1964.

Photograph

(occurrence from

15%

of cases) and secondary (occurrence from

in

5%) of foods
range. In
it

is

shown

Table 67 for seven different regions

in the

are placed only those foods

at least in

and groups were given which

were found,

one case, with an occurrence higher than 5%.

In the western part of the range, the

main food

is

shown

to

be

the water vole (Tatariya, western Siberia), and in the eastern

voles and mice. In the east, a

dents of moderate size

Daurian

number of main foods include roand Altai* pikas [Ochotona

daurica, O. alpina], Altai zokor [Myospalax myospalax], locally,

chipmunk and muskrat, and

also squirrel [Sciurus vulgaris] (espe-

cially during migration periods of the latter)

and jerboas. In

differ-

ent seasons, fish are of great importance in

*In the Russian original, this

some

places. In Ussuri
is

word

is

spelled "Alaiskaya", but this region

west

of the one being discussed, and "Altaiskaya" was probably meant

Sci. Ed.


1073

territory, carrion

marten

essential role in

food remains from wolf and yellow-throated are very important winter. Everywhere, birds play an a food of kolonok, chiefly small ones
in

to

lesser

on domestic birds are a widespread occurrence, as is damage to trapped hares, squirrels, kolonok and other animals. Feeding on reptiles and amphibians is observed at in Tatariya and especially in Primor'e, the periphery of the range Among a number of plant foods are found nut pine, "nutlets", fruits of actinidia vines and others. However, in the montane taiga
extent, poultry. Attacks

714 of the Altai, the

kolonok does not feed on nut pine "nutlets" at all (Ternovskii, 1956). In years with low numbers of mouse-like rodents, it suffers from hunger and eats carrion.

The dynamics of seasonal nutrition are known only for the mountains of Sikhote-Alin' and northeastern Altai; in other cases,
there are data only for the winter period. In Ussuri territory, the

main food throughout the whole year consists of mouse-like in Junerodents. Carrion is eaten from November to March; birds
August, and fish
715

in September-October (migration of salmonids).

The

role of plant food (not

above 4.9%) increases

in

August

February. Amphibians, reptiles and invertebrates are foods of the

warm
is

period of the year. The frequent occurrence of pikas (25%)

characteristic of the food of the Altai

kolonok

in the

summer

period.

The character of foods

varies quite noticeably not only in dif-

ferent geographical regions, but also in different habitats. In the

Sikhote-Alin mountains, the significance of carrion ranges from

to

20.8%, of birds from 5.4

etc.

to

12.4%, plant foods from 1.0 to

8.0%,

(food was studied in 12 habitats; Yu.A. Salmin and

to year. In

V.D. Shamykin).

The proportions of other foods change from year

Tatariya, the proportion of fish sharply increases in years of re-

duced occurrence of few water-and small voles (to 33%) (Grigor'ev and Teplov, 1939). The kolonok eats about 100-120 gm of food per day (Uspenskii, 1933). The kolonok usually makes a store of food, which sometimes happens to be quite significant up to 16 striped field mice [Apodemus agrarius] (Fetisov, 1936). Home range. Apparently, the home range of the kolonok is quite considerable, since it moves from 1.5-2 to 6-7 km per day,

and even up
to

to

8-10 km.

In the northeastern Altai, the area of

daily activity constitutes 1.5-2.0

km^ while

the

home range

is

equal

4-5 km^.


1074

Burrows and
varied.
It

shelters.
a

Shelter chosen by the kolonok

is

hollow in old fallen logs, empty stumps, piles of brushwood, space under the roots of fallen trees and under logs. The kolonok also often inhabits burrows of other animals (chipmunks, water voles, pikas and others), widening and deepening them. In western Trans-Baikaliya, the length of burrow pas-

may be

sages of kolonok appear to be from 0.6 to 4.2 m, and depth

0.2 to 1.3 m.

from

The nesting chamber

is

is

located in the middle or at

the end of the passage, and

lined with bird feathers or

wool of

mouse-like rodents (Fetisov, 1936).

Within the boundaries of
its

home

range, the adult kolonok has,

beside the permanent burrows, up to five temporary shelters situated

at a

distance of several kilometres from one another (Fetisov, 1936),

ity

Daily activity and behavior. Crepuscular and nocturnal activis typical of kolonok, but its rhythm is not clearly expressed
is

diurnal activity of kolonok

especially in

a quite
It

common phenomenon,
frost occurs at

summer and autumn.

hunts in the morning and

1956). During

during the day, as well as in winter

night and during the day

when severe

warms up (Shaposhnikov,

times of severe frost and snow-storms, the kolonok does not venture out of
its

snow

shelter for a series of days. In the forest-steppe

of western Siberia, this period of very low activity falls at the end of December-first half of January (Velizhanin, 1931).

The length of

its

daily tracks in winter in western Trans-

Baikaliya (Fetisov, 1936) ranges from 3-4 to 8 km; in western

Siberia and Kazakhstan (Sludskii, 1953)
in the

northeastern Altai

from

to 10

km

and more, and

Its

1.5-2.0 to 6-7 km.

length de-

pends on the quantity of prey and condition of snow cover (Shaposhnikov, 1956). In contrast to sable, in winter the kolonok under logs, in hunts food more actively, mainly under the snow bushes while sable snow-laden arches of beneath the windfalls and snow and frozen Compacted more often lies in wait like a cat.

crust

makes hunting under

it

the

snow almost impossible

for the

kolonok, and therefore,

confines itself to the northeastern slopes

of montane taiga in the Altai, and terraces of the northern quadrant,

where the snow

is

always loose and where frozen crust does

a distance of

not form (P.B. Yurgenson). During one night, the kolonok enters
into the

snow more than 30 times during

up

to 1.5

km. The kolonok goes up to 50 m under snow without exiting to the surface (Shaposhnikov, 1956). The kolonok rarely gets up into

1075

trees.

Following water voles,

it

swims

easily and quickly, going

out far from shore (Sludskii, 1953).

Seasonal migrations and transgressions. Seasonal migrations

(autumn-winter) from one habitat
716
to

another are

known

in

the

kolonok, as well as migrations for considerable distances related to

food insufficiency; however, factual data on these translocations

are insufficient since they

were often confounded with sharp

Fig. 256.

creek.

Tracks of a young male kolonok on soft first snow on the ice of a mountain "Kedrovaya Pad'" preserve, southern Primor'e. March 1964. Photograph

A.G. Pankrat'ev.

1076
fluctuations in numbers. Data on massive migrations of kolonok in

western Siberia and northern Kazakhstan reliable. The massive transfers were observed
tion,

in

1928 are more

in a northern direc-

and were associated with reproduction in water voles. In October 1928, migration of kolonok and ermines was observed through the city of Tomsk from west to east. Active broadening of
the range of kolonok
is

associated with years of significant in-

crease in its

number

as a result of particularly favorable feeding

conditions, and

may be

regarded as a particular type of migration.

in

Seasonal migrations occur

tions of food capture

connection with changes

in

condi-

and

their dispersion

among

habitats.
is

Reproduction. In the kolonok, the period of rut

tended.

quite ex-

The timing

is

subject to geographic variation: in western

Siberia, the signs of ruts appear at the beginning of February,

continuing through the end of March (Velizhanin, 1931; Zverev,

717

1931). In Primor'e, rut proceeds from the first third of

March and
to

continues to the end of April. In the Pushkin fur sovkhoz near

Moscow,
15

rut in six pairs of

kolonok took place from 25 April

May

(G. Uspenskii, 1933).

Mating lasts up to 35 min. and occurs repeatedly. Timing of pregnancy ranges from 38 to 41 days (Sludskii, 1953), and accordwithin limits of 35-42 days (Kler, 1941). Paring to other data

turition

was also observed after 28 days subsequent to the last mating, which is considered the minimal period of pregnancy (G. Uspenskii, 1933). The number of young (in caged conditions) ranges from 4 to 10 (7 on average). Growth, development and molt. Young are born blind and almost naked, with sparse white wool. After some days, they become covered with light yellow wool. Their eyes open on the 28th-30th day. Lactation stops at the end of the second month. By the end of August growth stops, and the young differ from adults only by
their darker color, the

deciduous tooth formula, and the less mas-

sive bones of the skeleton. Also at the end of August, the litter

begins to disperse.
718
in spring and autumn. a year Enemies, diseases, parasites, mortality, competitors, and population dynamics. Sable, both species of ferrets, red fox and eagle

Molt occurs twice

owl belong among the enemies of the kolonok. Ferrets, mink, sable, ermine, solongoi and red fox may be competitors of kolonok in the capture of food. Diseases have not been studied, but cases

1077

717

Fig. 257. Tracks of kolonok (left) and otter on snow along a montane creek in a place of typical habitat of kolonok. "Kedrovaya Pad' " preserve, southern Primor'e.

February 1965. Photograph A.G. Pankrat'ev.

of massive mortality of kolonok from

(Sludskii, 1963).

unknown reasons

are

known

Parasitic infection of kolonok, according to the data of Sikhote-

Alin'

considerable. In

(Yu.A. Salmin and V.D. Shamykin), is very summer, mites {Ixodes persulcatus) cling to them, and late in autumn Dermacentor canina. The frontal sinuses of all adult individuals are infected with the nematode Scriabingulus nasicola. Filaria is encountered in 69.4%, and ascarids in 24.4% (from 1015 to 30 individuals). Infection with other endoparasites is low.
preserve

1078
Fluctuations in numbers of the kolonok are

known

within the

borders of the whole range and are closely associated with the
yield of mouse-like rodents or water voles. Usually, years with an

numbers of kolonok follow years of massive rodent some regions of western Siberia, increases in numbers of kolonok were recorded in 1910, 1916-1917, 19221925 and 1928 (Lavrov, 1937). There is a basis to propose that years of intensive colonization of kolonok beyond the borders of its range coincide with years of increase in its numbers and the
increase in

reproduction. For

very colonization

is

associated with food deficiency in the regions

of reproduction (Lavrov, 1937; Sludskii, 1953).

Field characteristics. Tracks of kolonok on the snow resemble

those of ferret, which makes their observation difficult in regions

of co-occurrence. Paw prints of kolonok are smaller and more expanded than those of sable. The distance between the paired prints of the paws (length of leap) in an undisturbed running animal is 35 cm; a frightened kolonok makes leaps of up to 1 m. In contrast to ferrets, running of the kolonok has a somewhat confused character, is twisted, and more closely approaches that of mink, ermine and weasel (P.Yu.).

Practical Significance

The kolonok
is

is

a valuable fur animal, having a significant share of

fur preparation of Siberia

and the Far East. The fur of the kolonok

are highly valued: used in the prepara-

used as both natural and for imitating more valuable species.

hairs of the
tail

The guard

tion of brushes for painting. Settling near villages, the kolonok

may cause known harm

to poultry, but at the

same

time,

it

is

useful

for destroying rodents. In regions of intensive muskrat raising, the

kolonok may also be considered harmful. On the whole, however, the kolonok is undoubtedly a useful and valuable animal. The kolonok is hunted by shooting with dogs, or various box traps, (kulenki, plashki, cherkany) and rarely with jaw traps (P.Yu).

RUSSIAN, OR EUROPEAN, MINK

Mustela (Mustela) lutreola Linnaeus, 1761
1761. Viverra lutreola Linnaeus. Fauna Suecica,
1777. Lutra minor. Erxleben. Syst. Regni Anim.,
p.
1,

5. p.

Finland.

451. Re-

naming of lutreola Linnaeus.

1079

1879. Lutreola europaea Homeyer. Zoolog. Garten, 20,

p.

184.

Renaming of
1912.

lutreola Linnaeus.

Mustela

{Lutreola)

lutreola

wyborgensis

Matschie.
p.

Sitzungsber. Ges. naturforsch. Freunde Berlin. 1912,

347.

Vyborg, north of Leningrad. 1912. Mustela (Lutreola) lutreola cylipena. Matschie. Ibidem,

p.

348. "Gross-Shirau, vicinity of Valau, East Prussia" (be-

tween Chernyakhovsk and southeastern corner of Kursk Gulf,

Kaliningrad
719
district;

V.H.).
p.

1912. Mustela (Lutreola) lutreola budina. Matschie. Ibidem,

349. "Ortelsburg, East Prussia" (Shchitno southeast of

Ol'shtyn [Olsztyn], northern Poland; V.H.).

1912. Mustela (Lutreola) lutreola varina. Matschie. Ibidem,
Vic. of Shverin, Mecklenburg.
p.

351.

1912. Mustela (Lutreola) lutreola albica. Matschie. Ibidem,

Levitts [Lewitz] river

p.

351.

tributary of the Elbe.

Mecklenburg.
p.

1912. Mustela (Mustela) lutreola glogeri. Matschie. Ibidem,

354.

"Brig

[Brzeg],

Silesia"

(Bzheg, southeast of Vrotslav

[Wroclaw], Poland; V.H.).

1932. Mustela lutreola transsylvanica. Ehik. Allat. Kozlem, 29,
p.

142. "Komitat Turoch, Austro-Hungary" (district south of

Beskid,

Czechoslovakia Krakow; V.H.).

about

150

km
p.

southwest of
142. Kovachna;

1932. Mustela lutreola hungarica, Ehik. Ibidem,

Transylvania, slightly west of the sources of the Buzeu

[Buzaul] River, Romania, Nomen praeoccupatum Mustela eversmanni hungarica Ehik, 1928. 1939. Mustela (Lutreola) lutreola turovi. Kuznetzov. In: Novikov. European mink, p. 47. Caucasus. 1939. Lutreola lutreola borealis. Novikov. Ibidem, p. 63. Milet village, former Bogorodsk Co. Moscow governance. Nomen

praeoccupatum
1862.

Mustela flavigula

var.

borealis Radde,
63.

1939. Lutreola lutreola caucasica. Novikov. Ibidem,

p.

Prokhladnaya Station, northern Caucasus. Nomen praeoccupatum Mustela nivalis caucasicus Barrett-Hamilton,
1900.
1944. "L. (utreola)
I.

(utreola) turovi

Kuznetzov

et

Novicov."

Bobrinski. In Bobrinskii, Kuznetsov, Kuzyakin. Guide to

the

mammals

of

USSR,

p.

127. In place of Mustela (Mustela)

lutreola turovi Kuznetzov, 1939.

1080
1951. Mustela lutreola novikovi. Ellerman et Morrison-Scott.
Checklist Palaearct. Indian

Mammals,

p.

263.

Renaming of

Mustela (Lutreola) lutreola borealis Novikov, 1939.

1951. Mustela lutreola binominata. Ellerman et Morrison-Scott.

Ibidem,

p.

263.

Renaming of Lutreola

lutreola caucasica

Novikov, 1939 (V.H.).

Diagnosis
General coloration monotone and evenly brownish; venter same
color as dorsum or a bit lighter. Upper
lip,

lower

lip

and chin

white in color, sharply demarcated from dark color of head. White

spots of various size and form often on chest and along lower

weak its width equal to or Upper molar relatively small and longitudinal diameter of its inner lobe somewhat larger than outer part of tooth. Dimensions relatively large the condylobasal length of male skull more than 55 mm, of female, more than 52 mm (V.H.).
neck. Postorbital constriction of skull

more than width of

interorbital distance.

Description

Concerning general appearance, the mink is a typical representative of its genus and has a greatly elongated body with short limbs; however, compared to species described above, it is put together more compactly and does not look so elongated and thin. In this
respect,
it

differs

even from the kolonok and stands nearer

relatively quite large, with broad, but not

to

ferrets, greatly

reminiscent of the black (M. putorius).

tall,

Head

in

mink

ears (in winter, they slightly protrude

relatively well

from

fur).

Limbs

short,

developed membranes between

digits, particularly

720

on hind paws. These are larger than in ferrets and only terminal phalanx remains free. Tail relatively short, does not exceed half of body length, constituting about 40% of its length. It is covered by

more or
out,

less similar, close fitting hairs similar in length through-

and not fluffy the same type as other species of genus, strongly differing from fluffy tail of kolonok. Feet below weakly covered with fur. Digital and sole pads visible both in winter and summer; they have light color. Claws light-horn in color.

1081

Winter fur of mink very thick and dense, though not long, and

Compared to terrestrial species of the genus, underfur particularly dense. Guard hairs quite coarse, lustrous, and
quite loosely fitting.
as in all

"amphibious" mammals, contour hairs have very wide,

flattened middle part. Difference in length and density of fur

on

various parts of body, in particular on back and belly, relatively

very small. These and several other peculiarities of the mink fur
are associated with
its

semi-aquatic

way of
is

life.

Summer

fur

somewhat

shorter, coarser

and

less

dense than

winter one; however, this difference

small and incomparably

less than in other, purely terrestrial, species of the genus^^.

Color of winter fur

is

dark-brownish and of comparatively slight

variability. Particularly dark individuals are

dark-brown and even

almost blackish-brown in color, light ones have reddish-brown color. The tinge of reddish highlights varies, but is usually not great.

Color

is

evenly distributed over the whole body. In only a few

is

cases, ventral surface

a bit lighter than the dorsal side of the

720

Fig. 258.

Russian mink, Mustela (Mustela) lutreola L. Sketch by A.N. Komarov.

"Details on fur structure of Russian mink are absent in literature. American mink, acclimatized in our country has been very well studied by our authors. Data on this latter may also characterize the main peculiarities of our mink; however, it is necessary to keep in mind that the American mink possesses in the fur trade higher quality and is more highly valued; its fur is longer and more dense.

1082
body, and
in

particularly dark individuals and races, a dark, in

is

some even
Underfur
is

black, broad dorsal belt

is tail,

noticed.

Limbs

are slightly

darker than trunk, as

the end of which

may be

almost black.
is

bluish-gray.

Mink

fur, particularly

dark individuals,

very beautiful and has a dense silky luster due to dark guard hairs.

Mink has no
in

color pattern at

all

on the head, and

it

has a color

common

with the whole trunk, but upper and lower lips and

chin are pure white. Pattern of upper lip

chin, varies in size and form.
It

is quite stable, but on sometimes occupies the anterior part of the chin, and sometimes extends backward under the entire lower jaw; the posterior border is white, sometimes straight and

721

sometimes uneven. White markings sometimes occur on the lower surface of the neck and on the chest. They differ greatly in number, form and size. Sometimes, it is one small, hardly noticeable white dot, and sometimes, a sharply defined one, and sometimes several spots; in some mink, there is formed a long narrow white band going from throat to chest, in some cases, the chin spots extend backwards and

may be a spot or spots some places the whole rarely, in and not too on the chest. Finally, lower surface of the neck and the chest may be white. In this case, the spot is not smaller, and sometimes is greater, than the throat
occupies the throat region; with this there
spot of the stone marten. In individuals with a particularly large spot on the chest, white wool covering the digits of the fore- and
often also the hind feet, usually occurs.
In

mink with very dark

fur,

"graying"

may be

present

white

ends of individual guard hairs, or

throughout the pelage are

separate white spots,

some quantity of them spreading completely white. Extreme exotypic

deviations in the form of albino mink, or pelage covered with

may

occur.

Color of summer fur differs only slightly from winter. It is usually a somewhat lighter, dirty tone, with a more obvious
reddish highlights, with short, dull guard hairs
or silvery luster.

without dark

silky

On

the whole, individual variation in color, if white markings

on chest and throat are not considered, is not great. Sexual differences in color and fur character are absent. Young mink are clothed in dull, short and coarse "plump" fur. The first winter coat does not differ from the comparable coat of adults.

1083

Geographic variation
south.

is

relatively small (see below)

and

is

manifested in lighter and more reddish color of fur from north to

White markings on throat and chest are quite rare in the more frequent in a southerly direction; their general area increases and they attain their greatest development in the extreme south of the range (Caucasus). Just here and
north, and are encountered
relatively often

individuals with the described large white field

of "marten" type are observed, which are absent in the north, or

are represented there as completely individual rarities.

Compared

to the skull of the

kolonok, the skull of the mink

is

on the whole less elongated and with a less elongated braincase, with longer and more widely separated zygomatic arches, and with a relatively less massive and slightly longer facial portion. In several respects,
it

has characteristics similar to the skull of the black

ferret (M. putorius).

These apply mainly

to the relatively general

shortening of the skull, width of the zygomatic arches and several

other characters. Based on

some

characteristics, the

mink

skull

occupies an intermediate position between the skull of kolonok

and

that of the black ferret.

For the
characters.

rest, the
It is

mink

skull

is

characterized by the following

flattened, with a

moderately elongated braincase

and almost straight upper profile. Only in the region of the superorbital processes and somewhat behind them, it has a small
elevation; the profile of the facial part descends quite steeply
anteriorly.

The

postorbital constriction is
is

weak

its

width

at the

narrowest point
orbital width.

approximately equal to or greater than the interare relatively

The zygomatic arches

weak and

are

moderately widely separated, the mastoid processes are relatively

weakly developed. The auditory bullae are small, swollen, relaand narrow, and widely separated; in the posterior part, they separated from each other significantly farther than in
tively short

the anterior part.

The

ratio of their length to their

is

width

is

approxi-

mately

1.7.

Sculpturing of the skull

not sharp, crests and

is

prominences are moderately developed. The occipital crest

defined, the sagittal one
is

well

apparent, but quite weak.

The upper molar

is

small and

weak

its

area in projection

is

considerably less than the carnassial tooth and the longitudinal

diameter of the inner lobe

the second upper premolar

is

not greater or only slightly greater

than the longitudinal diameter of the outer.

is

The

posterior end of

in contact

with the anterior outer

1084

edge of the carnassial tooth, but

723
its

it

does not
the

anterior edge.

The inner surface of

lie in the groove on main apex of the lower

carnassial tooth

is

smooth, without a vertical edge.

On

the whole, appearance the specializations of the skull, as

a carnivore skull, are less developed than in ferrets

and even
is

less

than in the American mink.

Except for dimensions (see below) the female skull entiated by some less developed protuberances, crests,
lower weight, as
skull
is

differ-

etc.

and
its

in all other

members of
braincase.

the genus.

The juvenile

characterized by weaker development of the facial part,

shortness and a

more swollen

several of

Geographic variation of the skull, besides general dimensions, which exhibit growth from north to south, is manifested
extreme south (Caucasus) the elevation of the
its

in the fact that in the

skull in the interorbital region increases (strengthening

ity to the

similar-

kolonok) and several features appear (greater constric-

some of which make the Russian mink approximate to the American (M. vison; see below). The structure of the os penis is generally typical for the majority of species in genus, i.e. tip of the bone is bent upwards in the form of a hook and even somewhat backward, but the groove on
tion of the postorbital region),
is wider than that in the ferret and kolonok and the tip is bent in the form of a spoon (Novikov, 1956). If the bone is viewed from below, the terminal part deviates

the lower side of the bone

somewhat
to the

to the right.

to body and skull measurements, the mink belongs group of large species of the genus the ferrets and kolonok. Dimensions of the species as a whole: body length of males is

According

284-M373-430 mm, of females, 320-M352-400 mm; tail length of males is 124-M153-190 mm, of females, 130-M150-180 mm; length of hind foot of males is 54-M59-64 mm, of females, 50M52-53 mm; height of ear in males is 20-M23-25 mm. Male tail constitutes 35-M41-52% of body length, of females, 37-M4045%.
Condylobasal length of male skull
females, 52.8-M61.4-65.7
is

56.4-M62.7-68 mm, of
males
is

mm;

interorbital width of

11.0-

M13. 6-15.4 mm, of females, 11.1-M12.4-14.0 mm; zygomatic width of males is 30.8-M35.5-40.0 mm, of females, 28.9-M32.337.0

mm;

females, 26.5-M29.4-31.7

mastoid width of males is 27.0-M31.7-34.9 (Novikov, 1939).

mm,

of

mm

1085

;>1--. v^>~,y~..,

722

Fig. 259. Skull of Russian mink,

Mustela (Mustela) lutreola L.

1086

Dimensions of mink (18) from Volga-Kama Territory: body length of males is 310-M375-406 mm, of females, 300-M327360 mm; tail length of males is Ml 45 mm,* of females, 120-

M128-142 mm;
of females,
females,

length of hind foot of males

is

52-M56.2-61 mm,

45-M50.0-58 mm.

Condylobasal length of male skull is M67.5 0.40 mm, of M63.7 0.52 mm; zygomatic width of males is M35.2

0.52

mm,
is

of females,

males

12.9

0.30,

M30.5 0.65 mm; interorbital width of of females, 11.3 0.32 (V.A. Popov, 1949;

here also other measurements).

Length of OS penis of adult males (6) is 34.8-M35.9-37.8 mm, M33. 5-34.8 mm; its weight in adult males is 0.230-M0.292-0.380 gm, in young, 0.100-MO. 118-0.130 gm (V.A.
of young, (10) 32.6,

Popov, 1949). The total weight (from data for Middle Europe)

is

550

to

800

gm

(V.H.).

Systematic Position^^

The systematic
ferently

position of

mink among those species which

are

here united in the genus Mustela, has been evaluated entirely dif-

by different authors. An extreme point of view holds that European mink together with the American mink (M. vison) belong in a special genus Lutreola, usually standing as more or less closely related to the kolonok CKolonocus") and to polecats CPutorius"), more often to the latter. In such an evaluation of the characteristics of the mink, it is impossible not to see extreme
the

over-estimation, of a
724

known hypothesis

of their unique

way

of

life.

Right now,

this

point of view has almost no supporters.

More
genus

often, both species

were included, and are included,

in the

Mustela in the category of a separate subgenus, equivalent to ferrets.

Isolated opinion holds that minks represent a special subgenus

genus of polecats (Putorius; Stroganov, 1962). At the same time, in morphological features, mink have no characteristics which should force any higher evaluation of them. According to the sum total of characteristics, the mink itself represents a development and combination of the same set of features which characterize the previously described species of the genus.
in the

*No range
carnivores".

given

Sci. Ed.
in

"For section on American mink, see end of book

"Supplement

to the order of

1087

Within the limits of the genus,

it

is

most correct

to consider

it

close to the kolonok (M. sibirica) on one side, and on the other,

already displaying several characteristics of the ferrets, namely the

black M. putorius.

To

a certain extent,

its

position

is

intermediate,

connecting the group of true ferrets

{''Putorius''')

with the other

members of

the genus. Therefore, consideration of ferrets as an independent genus has become practically unsupportable.

The "intermediate"
that in nature hybrids
tered,

position of mink is emphasized by the fact between mink and black ferret are encounrarities.

and these do not represent exceptional

in

There are

M[oscow] U[niversity]) and they are described (Ognev, 1931). They are well known to furriers, who even have a special name for them ("khor'tumak") and affirm that they are met with "not rarely". On the
such specimens
(Z[oological]
of]

museums

M[useum

skins of hybrids, characteristics of one or the other parental species

crosses between

predominate (Kuznetsov, 1952). Remarks stating that "data on mink and ferret are completely absent (Novikov,
is

1939)

based on misunderstanding. Nothing

of "khor'-tumak".

is

known concerning

fertility

Of all species of the genus, the European mink is closest to the American mink (M. vison). This latter itself represents a further development of the mink type and compared to it, the European mink carries certain infantile features (larger protuberances, crests, etc. of the skull, development of postorbital constriction in the American mink and some other features see description of this

species).

Despite closeness in their general form, however, they cannot be considered races of one species, as several European authors have done (Zimmermann, 1959; Gaffrey, 1961). The morphological characteristics

tion of both species in nature,

which divide them are substantial. Hybridizawhich is sometimes assumed for

is

Central Europe (Gaffrey, 1961),

in

not confirmed.

On

the contrary,

our country a clear antagonistic relationship between the two

species in nature, and destruction of one by the other, has been

established (V. Popov, 1949, see also section on

American mink).

Attempts
is

to cross

both species in captivity were not successful.

In the sense of the structure and phylogenesis of the genus, it very interesting that the European and American minks, in

their degree of specialization

and development of separate con-

crete characters (postorbital constriction, for example), themselves


1088
represent species parallel to the black (M. putorius) and white (M.

eversmanni) ferrets (see also section on American mink) (V.H.).

Geographic Distribution
Europe except the south and northwest Caucasus, western part of
western Siberia.

Geographic Range

in the Soviet

Union

This constitutes the greatest part of the species range

time, nearly the whole.
It

at the

present

occupies the European part of the Union,

the range in the extreme northwest

Caucasus and the western part of western Siberia.

The northern border of

in Karelia, passes a little north

of the

extremity of Topozera [lake] and

725

Kem' Kuzema

river

at the

southern

[city]

(Marvin, 1950,

1951) or a little to the north (Kesten'ga), but it does not reach Kovdozero, i.e. it generally lies between 65" and 66 N or along 66 N. lat. References to its occurrence near Kandalaksha and even

Umba

(Ognev, 1931) and, in general, on the Kola Peninsula are mistaken (Pleske, 1886; Novikov, 1939). Farther, the border passes
along the shore of the White Sea to Arkhangel'sk, and thence along the sea coast or near
it

to the

mouth of

the Kuloi and

Mezen'. From here, the border

tundra, adjacent to

line,

passing south around the Kanin

Cheshsk

inlet

passes around Sula

(left tributary

and Malozemel'skaya tundra, of lower Pechora) from the north,

i.e.

and along the Pechora, extends as a projection, to the delta, 68 N. lat. (Novikov, 1939; V.Ya. Parovshchikov). This
northernmost point of the range.

to

is

the

Farther to the east, the border includes the entire lower and middle course of the Pechora, skirting the Bolshezemel'skaya tundra from the south along the crooked-forest belt, north around the basin of Usa (apparently, only the lower reaches of its right tributaries, Laya and Kolva), and ascends along its tributary, Ad'zva, to Khased-Khard. To the east of the White Sea, the northern bor-

der of the

mink range extends,

its

therefore, to the southern part of

the forest-tundra or along

southern boundary, moving up the

river valleys to the north. Locally, the

mink occurs only

as a tran-

sient (perhaps also living there) at the border of the true tundra
(at

Nes' village on the western bank of the Kanin, north of the

Arctic Circle).

1089
In the Urals, the location of the border is unclear. Apparently, descending from Khased-Khard directly, or almost directly, to the

south,

it

crosses the Urals

it^'.

somewhere around
mainly
at

the Arctic Circle or

probably south of

Farther, the border goes directly south, inriver (noted

cluding the Lyapin

Saranpaul') and the

in

upper course of Severnaya Sos'va. Thence,

an as yet undeter-

and then below Tobol'sk. From this point, the range border passes along the Ob' or a little to the north to the lower course of the Agan river, the right tributary of the Ob' (I. Laptev; perhaps transients). Thence, the border
turns eastwards reaching the

mined

path, directs itself, apparently, again southwards

Ob' somewhere a

little

line
is

goes directly south, covering the basin of the Dem'yanka (mink known from several places in its middle course) and reaches

on the Irtysh. Beyond the Irtysh, the range does not extend above its course (Yanushevich and Arkhangel'skii, 1952; I. Laptev,
1958).

western Siberia
to

The southeastern and southern borders of the mink range in is poorly known. On the whole, from it arches the Ural ranges, somewhere approximately at the latitude of
the border, apparently, passes

Chelyabinsk. South-southwest of
a
little

mink are known at Lake Chagly north of Kokchetav; Sludskii, 1953) and at Zverinoglovskaya on the Tobol south of Kurgan (I. Laptev, 1958).
north of Petropavlovsk (transient
Farther, the border follows along the eastern slopes of the

Urals southwards through Orsk to Aktyubinsk, then westwards along

the Ilek;
it

includes the lower course of the Utva (Chingurlau) and

at Ural'sk or slightly to its south. Mink do not occur lower along the Ural [river]. The suggestion (Novikov, 1939)

reaches the Ural

that

mink

is

absent in the southern Urals and inhabits only the

this region,

western slope in

Danilov, 1951; Kirikov, 1952). Probably transient mink are

at Ilek,

on the

Irgiz river

mation concerning the occurrence of mink in Alakor(Semirech'e; Shnitnikov, 1936) are lacking any basis.

(
is

is

incorrect (Shvarts, Pavlinin and

known

settlements; Sludskii, 1953). Infor-

From Ural'sk
the

the border, going west, extends over the basin of

Volga

tributaries

Bol'shoi
mink

Irgiz (Elpat'evskii, Larina

and

Golikova, 1950; mink are absent on the Uzen') and descending

"The
or
it

reference that the

distributed in the Urals nearly to 70 N.

lat.

(Shvarts, Pavlinin and Danilov, 1951)

is

a misprint; 70

is

was based on some sort of misunderstanding, the latitude of Vaigach Island.

1090
southward,
it

passes around the basin of the Eruslan from the east

(Orlov and Kaizer, 1933), and then reaches the Volga. Along the valley of this river, the range reaches the [Caspian] Sea as a nar-

727

row band. Somewhere in the region of maximum proximity of the Volga and the Don, the [range] border crosses over to the Don and descends to its mouth. Thence, it again passes at some distance from the shore along the Azov Sea approximately to Osipenko
(Berdyansk) and then, passing around the southern Tavrichesk steppes from the north, reaches the mouth of the Dnepr and along the shore of the Black Sea, passes beyond the frontier of our
country.

According to some data (Novikov, 1939), the border envelops the Donets ridge from the north. This is quite possible for the present time, but, apparently, the absence of mink in this region itself constitutes a secondary phenomenon. Equally with this, there are references about occurrence of mink throughout the Ukraine
(Migulin, 1927; Korneev, 1952; Sokur, 1960).
In the west, where the range reaches the state frontier, the mink mainly exists in the Trans-Carpathians (Konyukhovich, 1953). The mink is absent in the Crimea. The outline of the range of mink in the Cis-Caucasus and in the Caucasus is very complicated. The northern border of the Caucasian part of the range begins at the Don at Tsimlyanskaya

and descends southward

Thence,
it

to the Sal,

reaching

its

lower course.

passes to the Manych, extending along the river east-

wards

to the

mouth of

the Egorlyk (Veselovsk reservoir) or

some-

what farther, descending along Egorlyk southward and reaching the Kalaus a little above Ipatova. From this point, the border line
crosses over the
taries.

Kuma

basin, also including

its

northernmost tribu-

Along

the
it.

Kuma,

the range border runs to the sea, or some-

what short of

The southern border begins at the Caspian Sea somewhat south of Makhachkla (at Deshlagar) and directs itself northward along
the foothills, and then westward reaching Khasav'yurt. Thence,

along the base of the mountain and along the foothills, it goes to Ordzhonikidze (former Vladikavkas). Farther to the west, it ascends high into the mountains (Beskes, tributary of the Bol'shoi

Laba; slightly below Klukhor

in

at the

source of the Kuban),

it

goes
the

an irregular line through the upper reaches and sources of the

Terek,

Kuma

and Kuban river systems

approximately

to

u<

'^

1092

Tuanse meridian. Here, the border crosses the Caucasian range, and the range extends as a narrow strip along the southern slope to the southeast, reaching or almost reaching Sukhumi. Here, the border ascends quite high into the mountains (Pskhu, on the Bzyb'
[river]).

In the west, the

mink

is

distributed along the shores of the

in

Black and Azov seas (present

Taman'). In the
is

east,

between

Deshlagar and the mouth of the Terek, the range

limited to the

Caspian shore. In the semidesert, in expanses devoid of rivers between the lower Terek and Kuma (Nogaisk and Karanogaisk steppes), the mink is absent, and the eastern border of the range here goes along the periphery of the Kuma and Terek basins^^. As is seen in the description given the range of the mink in the Caucasus, despite the widely propagated view (Novikov, 1939, 1956), is not cut from the main region of occurrence of the species in the

European part of the USSR. In the west, between Karelia and the Black Sea, the range of the mink everywhere goes beyond our state frontier. There is a widely distributed view that the mink is related to a number of our mammals, the ranges which are expanding to the
east. It is usually

believed that the above-outlined region of occur-

rence in the Urals and beyond the Urals, was formed in the last

80-100 years, and in the 18th and first half of the 19th century, mink were absent in Siberia. It did not apparently, occupy the Urals, being met with only on its western side. The mink crossed
the Urals, apparently, only in the 70's of the present century and

attained the extreme eastern limits of

its

range only

at the

begin-

ning of the 30's of our century (by 1932).

It is

necessary, however, to keep in view that this opinion about

the colonization of

mink

is

based to a considerable degree on infor-

mation from Sabaneev

speaking,
little is

who is known for his ruinous errors. Generally known on the course of colonization in the mink
I.

(for details, see

Novikov, 1939;

Laptev, 1958). That

is

it is

conis

tinuing to settle
728

new

places at the present

not excluded. This

shown by
and the

the above-mentioned transgressions of

Irgiz.

Beyond

the Irtysh (upper

),

was

still

absent in 1950 and 1960 (for details,

Kokchetav however, the mink see Novikov, 1939).

to

mink

'Range in the Caucasus after Dinnik, 1914; Satunin, 1915; Turov, 1926, 1928,
1931; L. Boehm, 1929; Heptner, 1936; Heptner and Formozov, 1941; Vereshchagin, 1947; 1959 and other sources, and after material of V.G. Heptner.

1093

Equally with
in

this,

there

was

a suggestion recently, not lacking

serious foundation, speculating that

mink

in

western Siberia
in

occurred very long ago

even before the

academic expedition

the 18th century, but was always rare; and is rare now almost everywhere in Siberia (I. Laptev, 1958). It is possible that, with

colonization of

western Siberia, there is a situation analogous to that previously mentioned for the pine marten (see p. 844)*.
in

mink

*<:

?
0]

zr

Fig. 261. Species range of the Russian mink,

Mustela (Mustela) lutreola L.

is

In

Europe,

beyond

the borders of the

USSR,

the range

as established in the

19th beginning

of 20th centuries (after Novikov, 1939, with modifications).

*Page reference omitted

in

Russian original

Sci. Ed.


1094

On

the whole, factual materials supportive of this and other view-

points are scanty, and the question requires further study.

Information
directions.

is

absent concerning colonization of

in range are to

mink

in other

Some changes

be expected in connecin

tion with the introduction of

American mink

many

parts of our

country.

Geographic Range outside the Soviet Union

This (as established in the 19th and 20th centuries) includes Finland northwards to 66 N. lat. or a little farther, Poland, Hungary,
729 Czechoslovakia,

Romania, [eastern]* Austria, Bulgaria north of

i.e.

the

Danube and Dobruja**, northern and western Yugoslavia

in the
lat.

Croatia, Bosnia and Herzegovina countries south to approximately 43 N.

Istria,

west of those and in the east, to Belgrade, the territory of the German Democratic Republic, a considerable part of France (except the southwest and northeast ***). The mink is absent in West Germany, England, Ireland, Denmark, the Scandinavian Peninsula, [western]* Austria, and Switzerland

and the remaining part of Europe^^.

The mink

is

rapidly disappearing in the west

its

beyond the boundhas already disap-

ary of our country and

range

is

shrinking.

It

peared or almost disappeared from Central Europe where it still lived in the beginning of the 20th century. One of the causes of the
disappearance of the European mink
lies chiefly in its destruction

in

by the

western Europe apparently American [mink] (V.H.).

Geographic Variation

quite large

number of

races have been described from the

if

comparatively small species range of the mink, even

Russian
original

we do
is

not

ambiguous

Sci. Ed.

*Not

clear; Bulgaria lies entirely south of the

Danube, and Dobruja

a region

of northeastern Bulgaria and adjacent Romania Sci. Ed. ***In Russian original, "northwest", a lapsus Sci. Ed.

''Reference to Italy (Bobrinskii, 1944)

may

apply only to the Istrian Peninsula,

within the boundary of Yugoslavia. In Italy proper, mink is absent and was also absent in the period under review. Data concerning Bulgaria and a portion of Yugoslavia were given by B.M. Petrov. References encountered in the literature concerning

now

mink in the Scandinavian Peninsula and its disappearance replacement by the American mink (crowding it out or its absorption by hybridization), are incorrect. It was never present there (see Ekman,
the previous presence of

not too long ago and

1922).

its

1095
take into consideration the notorious

synonyms given by Matschie

first

(1912). In actuality, geographic variation in the species,

in the

of

all

USSR, where

the largest part of

its

range

lies, is insignifi-

cant and less than

is usually reputed to be. This applies both to the amplitude of variation of characters and to the number of races.

The

relatively small geographic variation in animals associated

is

with water

entirely natural. Usually, six or seven races are

recognized, of which there are 5 or 6 in the

USSR

(Novikov, 1939,

1956; Ellermann and Morrison-Scott, 1951). Such a large quantity

partially

Scott, 1951, of the

comes from misunderstanding (Ellerman and MorrisonCaucasian form, for which two names and two

forms are given).

General dimensions, color, density and fluffiness of the fur are
subjected to geographic variation. These characteristics change
regularly from north to south

dimensions
On

increase, fur lightens,

coarsens and becomes less dense.

acteristics,

the whole, both these char-

and also

in the distribution of the

geographic forms of

mink, require solid revision.

In our country,
it

is

quite evident that

two or three forms can

be distinguished. It is possible that several others, highly doubtful forms are still present (chiefly after Novikov, 1939 and Kuznetsov, 1941, 1952 with modifications and additions after materials of the
Z[oological]
1.

M[useum

of]

M[oscow]
/.

U[niversity]).

Northern mink, M. (M.)

lutreola Linnaeus,

1761 (syn.

minor, wyborgensis).

Color dark, brownish-chestnut colored or even dark-brown with

diffuse broad belt on back. Tail tip black. Underfur dark, bluishgray. Pelage long,

compact and
is

silky.

Body
length
is

length of adult males

usually

124-M 137-1 50 mm;

length
is is

length of hind foot

340-M365-380 mm; tail is 54-M56-58

of body length. Weight

mm;

tail

36-42*, averaging

38%
is

of 100 skins

6.5 kg.

Condylobasal length of male skull

females, 52. 8-M58. 2-64.3

56.4-M61. 5-64.3 mm, of

males
is

mm;

interorbital width of

11.0-

M13.4-15.0 mm, of females, 11.1-M12. 8-14.0 mm; zygomatic width of males is 32.4-M34. 1-36.9 mm, of females, 28.9-M32.137.0 mm; mastoid width of males is 27.0-M29.7-32.2 mm, of females, 26.5-M28.2-31.7 mm.
*In Russian original, mistakenly given as

"mm",

rather than

% Sci.

Ed.

1096

Found

in

the northern

European

part of the country south-

wards approximately to the line Leningrad-Yaroslavl'-Gor'kiiKazan'-Sverdlovsk; possibly

Outside the
730
2.

in Bashkiriya.

USSR in

Finland.
/.

Middle Russian mink, M. (M.)

skull

novikovi Ellermann et

Morrison-Scott, 1951 (syn. borealis).

Body and

dimensions moderate
in

somewhat larger than

preceding form.

Color somewhat lighter than

nominal form, dark-tawny or

dark-brown, usually with light film of reddish highlights. Dark belt on the back weakly defined or absent. Underfur lighter than in northern mink. Pelage dense and compact, but shorter and less
dense, and considerably less silky than in preceding race.

Body

length of adult males

is

360-420 mm. Weight of 100

skins about 8 kg.

Condylobasal length of male skull is 59.2-M63.7-66.8 mm, of females, 57. 2-M58. 3-59.4 mm; interorbital width of males is 11.9M13. 8-14.7 mm, of females, 11.3-M12. 3-13.9 mm; zygomatic
width of males
36.7
is

30.8-M36.3-38.6 mm, of females, 31.2-M33.0is

mm;

mastoid width of males

28.4-M3 1.2-33.2 mm, of

females, 28.0-M29.2-30.3

mm.
form
in west,

Found

in

middle zone of European part of Union, south of

including Estonia, east-

distribution area of previous

ern part of Latvia, Byelorussia, southward to border of species, except the Cis-Caucasus and Caucasus, and, probably, eastern Ukraine, Lower Don and Lower Volga regions. The position of

mink
3.

in

Bashkiriya

is

not clear.
not encountered.

Outside the

USSR, probably

Caucasian mink, M. (M.) I. turovi Kusnetzov [in Novikov]*, 1939 (syn. caucasica, binominata) Body and skull dimensions large the largest of the races living within the boundary of the USSR. Skull relatively massive

with well developed protuberances, crests, etc. and strong zygomatic arches. Postorbital constriction strongly expressed. Line of upper
skull profile in region of supraorbital processes

somewhat

elevated.

Pelage quite long, but sparse and rough, with less compact underfur. Color light-tawny or light-brown with clear rusty (reddish) highlights. Underfur light, bluish-gray. White markings on
chest frequent, and
*See note below

much more

often than in other forms; in

many

Sci. Ed.

1097
cases, large white spot located on chest,

sometimes covering entire

throat and anterior part of chest. Often, in such mink, ends of

limbs also white.

Body

length of males usually

more than 42 cm. Weight of 100

is

skins about 9 kg.

Condylobasal length of male skull

of females, 54. 0-M56. 3-59.4

60.8*-M58.2-68.0 mm,
males
is

mm;

interorbital width of

12.7-M14.0-14.9 mm, of females, 12.0-M12.5-13.2 mm; zygomatic width of males is 34.0-M36.8-40.0 mm, of females, 32.0-M33.335.8 mm; mastoid width of males is 29.4-M3 1.8-34. 9 mm, of
females, 30.5-M30.8-31.2
In

mm.

Caucasus, Lower Volga and Lower Don regions; probably

eastern Ukraine.

Absent outside USSR.

Slight elevation of skull in region of supraorbital processes

resembles somewhat the corresponding structural peculiarity of kolonok. On the other side, dimensions and relatively massive skull

and the more pronounced postorbital constriction combine to make skull of Caucasian mink somewhat similar to American mink skull. Note. The name turovi, given to Caucasian mink, appeared in literature in 1939 in the work of G.A. Novikov, based on B.A. Kuznetsov's manuscript on page 47. On page 62, G.A. Novikov
describes this same Caucasian

mink

as a special

new subspecies

under the name caucasica, putting the name turovi of Kuznetsov

731

synonym. Later, N.A. Bobrinskii (1944) applied the name "turovi Kuznetzov et Novikov, 1939" to the Caucasian mink. In this draft, the name conformed and was used later (Novikov, 1956). What G.A. Novikov (1939) placed in quotation marks on page 47 was a complete diagnosis from B.A. Kuznetsov's manuscript with his data showing distribution, more precise type locality and occurrence (see above, synonymy of species); the Caucasian form of the mink must be named as given above, but not attributed to G.A.
as
its

Novikov or

to

both authors.
is

Of
They

the three described forms, the most distinguishable

the

northern,

M. (M.)

I.

lutreola and the Caucasian,

M. (M.)

/.

turovi.

are well delineated forms differentiated both by their body and skull dimensions, and also fur color and characteristics, and by

*Sic\ judging
Sci. Ed.

from values of mean and maximum, probably should be 50.8

mm

1098

development of white markings on the chest. At the same time, all these features are more or less steadily deployed from north to south and there are no sharp limits between [southern and] polar form. The Middle Russian form, M. (M.) I. novikovi is actually a transitional population between M. (M.) /. lutreola and M. (M.) I. turovi. On the whole, it is closer to the northern form and with this prior information, it might have been united with it. At the same time, the differences of the Caucasian form remain sufficiently evident. Therefore, in the European part of the USSR, from north to south, two forms may be recognized. Fur standards relate the Middle Russian mink, with some reservations, to the northern sort, which acknowledge two in all. Nothing is known about the systematic characteristics of the Siberian mink. No differentiation from the described forms should be presumed to have occurred. 4. Middle European mink, M. (M.) I. cylipena Matschie (syn.
budina, varina, albica, glogeri, hungaricaY^.

Dimensions very large, only slightly inferior to Caucasian mink, turovi. Color quite dark, apparently, corresponding to I. color of Middle Russian mink M. (M.) /. novikovi. Body length of males is 420-430 mm, of females, 370-400 mm; tail length of males is 160 mm, of females, 140-180 mm. Basal skull length of males is 58. 8-M60. 6-63.1 mm, of females, 53.3-M54.0-54.9 mm; interorbital width of males is 13.3M14.2-15.2 mm, of females, 11.8-12.0 mm; zygomatic width of males is 35. 5-M37. 3-38.4 mm, of females, 32.4-32.6 mm; mastoid width of males is 30. 8-M3 1.9-33.3 mm. In Kaliningrad district, Lithuania, and western Latvia. Outside the USSR in Middle Europe, except the extreme west (France) and, probably, in Hungary, Romania and Yugoslavia:

M. (M.)

Poland.
very little known, essentially doubtful form, the characterand distribution of which are poorly studied in our country and which cannot be considered established; the same, it can be
istics

said emphatically, also applies to Central Europe, where, more-

over, the

mink

is

exterminated or almost exterminated

at the

present

time. Establishment of the actual nature and independence of

this race is
""The

only possible
is

now on
it

the basis of material from our

name hungarica
(see

considered to be a
is

synonym of

the

form transsylvanica.

As shown above
type locality

synonymy), of the form gloyeri,

described from the immediate vicinity of the

cylipena.

i.e.

1099

excluded that the Middle European form is very and may be identical to, the Middle Russian. 5*. Romanian mink, M. (M.) I. trans sylvanica Ehik, 1932. Dimensions smaller than those of Caucasian form. Color darktawny. Found in Moldavia (?) and Carpathia.'*'
territory. It is not

close,

Outside the

USSR in

Rumania, possibly Hungary, Bulgaria

and Yugoslavia.
732

To Of

the highest degree, a poorly

known and

entirely doubtful form.

the described forms having a relationship to our fauna,

only two or three (nominal, Caucasian and Middle Russian)

basic study

may

actually be considered real; the remainder are very doubtful and

is

required of geographic variation of the species in

the Pribaltik, the western parts of the Ukraine, Poland and

Middle

and southeastern Europe.

Outside the borders of

recognized, M. (M.)
/.

USSR,

still

another form

is

usually

biedermanni Matschie, 1912

Biology

France (V.H.).
range has the
is

Population. The distribution of the mink within

character of a fine lacey network, since
to a
its

its

innate area

restricted

narrow ribbon along the banks of

its

preferred small bodies

of water. Therefore, density of settlement in different regions

depends on frequency of water bodies within the area and the

degree of their suitability for the species.
ent the density of

On

the latter

is

depend-

mink within

the

home

range.

Mink populations and distribution are difficult to determine. Although forests are not absolutely necessary for its occurrence, they nevertheless create the most favorable conditions. Therefore, the majority of mink inhabits the forest zone, and a very small number inhabits the forest-steppe, and even fewer, the steppe zones. Thus, in the northern zone of the European part of the
*6
in

Russian original
to

Sci.

ed.

"According

some data (Konyukhovich, 1953),

district),
is

the mink,

which

is

quite

widely distributed in Carpathia (Zakarpatsk

relatively large

and differs from the Middle Russian form. It is now impossible to these data whether it is the given form or the Middle European cylipena.

and dark decide from

1100
country,
districts

24%
and

of mink were captured, in Arkhangel'sk and Vologod

in

and Urals
If,

23%,

Komi ASSR
and
in

25%,

in the

Volga

area, Bashkiriya

Kareliya and Leningrad district

15%.

in the recent past, the

number of Russian mink

skins taken

annually was equal to 50,000-60,000 (V. Popov, 1964), then at the

beginning of the 60' s,

trapping industry,

it became two three times less. The causes of this decrease are several: a general decrease in intensity of the

crowding out from a series of areas by the numbers due to increasingly unfavorable living conditions etc. The replacement of natural mink fur by cage-bred American mink in the world fur market played a known role. At the present time (mid-60's), judging by the catch, the highest populations of mink are found in Kostromsk, Yaroslav, Kirov, Gor'kii and Arkhangel'sk districts (about 25% of total catch in USSR); i.e., mainly in the southern taiga zone.
its

American mink,

local reduction in

Habitat.

Most

typical for

mink

are small bodies of fresh run-

ning water.

It is

rarely encountered on large rivers; mainly in flow-

ing valley lakes and at mouths of tributaries.

The

riparian zone of

large and moderate rivers are considerably inferior to the small

creeks with

tective conditions. Nevertheless,

as

abundance and the promet with on such rivers the Volga, Kama, and Vyatka (V. Popov). Mink has not masas regards food

woody banks

mink

are

tered the water surface

it

is

an inhabitant of the riparian zone.

the water in the

The dimensions and depth of

summer

period

do not have primary significance for the mink. Only in the region of the brood den is a depth of not less than 0.5 m necessary for masking the underwater entrance. In winter period, the presence and abundance of unfrozen sections

polynyas, broken

ice, springs

are

very important places.

at a

No
ice.

less important is freezing of the

it

water

high level, with

formation then, when

winter food

recedes, of a considerable space below the

is

Total freezing of the water

is

very unfavorable, but

when

abundant or when accumulations of hibernating frogs

is

are available, winter survival

not inhibited.
are

Water bodies where "naled"

surface] are unsuitable for mink.
feel
733

formed [water on top of ice With incomplete freezing, mink

winter in very small bodies

themself

in excellent conditions in

wide and not more than 1 m deep. In one place, mink prefer steep banks and in another low, marshy ones. The security of riparian zone and channels litter, availability of
of water

1.0-1.5

1101

tree

and shrub vegetation or bogs with timber along the bank are

entirely essential for the well-being of this animal.

Water bodies colonized by mink

of water bodies as denning areas,

are extremely variable as re-

gards feed conditions, denning and protection. All of the variation

may be grouped

into seven types:

1) small water bodies with semiaquatic vegetation and low, marshy banks; 2) small water bodies of the meadow type with low marshy

banks bordered with willow herb and speckled alder; 3) small meadow water bodies with steep banks, with an interrupted border of speckled alder; 4) small water bodies in coniferous and mixed forests; 5) small water bodies with water meadow of the broad-leaf
type; 6) black alder

swamps;

7) montane-taiga creeks

and rivulets

with rapid current and rocky channels.

Mink

often settle at mill

dams,

in flood lakes, tributaries

and dry river beds, sometimes also

in floodlands

of large rivers. Winter regimes of water bodies most

sharply restrict the distribution of mink.

Food. In composition the food of mink includes almost

elements of the aquatic and riparian fauna.
Its

all

food varies depend-

ing on composition of this fauna, time of the year, crops of different foods and changing conditions of their availability.

The foods of European mink are poorly studied. Voles are the most significant (36% of occurrences), fish (28.8%), crustaceans (26.7%), frogs (17%) and water insects (19.8%). In Tatariya (floodlands of the Kama), fish is the most important (35%), while mammals, (29.5%), amphibians (18.1%), birds (4.5%) and insects (3.4%) are less so. There are no essential differences between its food and that of American mink (Grigor'ev and Teplov, 1939). In individual cases, mink is capable of overtaking and catching any fish weighing up to 1-1.2 kg, but it usually prefers smaller ones. Often but not everywhere, mink feed on crustaceans. They
are absent not only in the Trans-Urals but also in

many water

bodies

European parts of the Union. The daily food requirement of the mink food consists of 140180 gm. When food is abundant, mink makes stores. Usually, they are not great and contain, for example, up to 20 loaches
in the

[Miagurnus], 3 large ide and pike or several water voles. Sometimes,

the store consists of frogs alone.
kills,

When

there are outbreaks of fish

the store of

mink

increases to 10-15 kg (V. Popov, 1941).

Local fluctuations in the abundance and conditions of capture

of different foods always occur everywhere, and these are reflected

1102
in

mink

nutrition. Thus, for

example,

in Tatariya

(Grigor'ev et

al.,

1931) in the winter of 1929/30 the frequency of occurrence of

On

water voles was twice that of the preceding year (14.2 and 33.3%). the other hand, the frequency of occurrence of frogs in its food

decreased more than two times (11.1 and 27.7%), of fish

by
is

al-

most 2.5 times (27.7 and 67.8%), and birds completely disappeared from
its

ration. This is explained

by the

fact that water voles

were
not

plentiful

and

this

food

is

more

calorific than fish

and frogs.

Home

range. The range inhabited by individual animals

food resources on the banks of small water bodies. The dimensions of individual home
large, possibly thanks to the generally high

ranges fluctuate depending on feed resources of a given region. In

regions of water

meadows with

little

food, the

home range

attains

734

60-100 ha, but significantly more often, the mink lives in an area from 12-14, and up to 27 ha. In summer, the range is smaller than in winter. Along the shoreline, the length of the home range fluctuates from 250 to 2000 m, with a width of 50-60 up to 100 m. The length of daily movements of mink differs. In spring, individual wandering males move from 4-5 up to 7 km per day. In autumn from 50 to 1000 m, sometimes the whole movement is situated within the boundary of a water meadow not more than 100 m long (L.G. Kaplanov). In winter, its movement on the snow surface decreases to 100-150 m per day. In the Moscow district in the 30' s with very intensive commercial harvest the catch of mink per 1000 ha of denning area consisted of 50 to 200 individuals

(P.B. Yurgenson).

Burrows and

shelters.

Mink

has both permanent burrows, as

well as repeatedly visited temporary shelters.

freshets and

The burrow

is

used

throughout the year, with the exception of the period of the spring
floods. Usually, the burrow is located not from the water's edge. It is of simple construction: 1-2 passages 8-10 cm in diameter and 1.40-1.50 m in length, leading to a nest chamber of dimensions approximating 48 x 55 cm (Flerov, 1926). In the brood burrows, this chamber is usually lined with dry grass, moss, mouse wool or bird feathers; some-

summer

more than 6-10

times, a lining does not occur at

all.

Often, the burrow opens to the outside under water, as in the

and desman. Such underwater outlets are conbank of the water body, though not high, is abrupt and steep. In other cases, the burrow is made under the
river beaver, otter

structed

when

the

1103
roots of trees, while in

marshy places, the burrow

is

made

in

high
set-

hummocks
tle in

"kobla" of European alder or spruce. Often, mink

on the banks or in hollows in the wind-felled branches and fallen trees. Sometimes, it constructs a nest in heaps of reeds, brushwood or under the protection of overhanging sod and woody roots of a steeply rising bank. Near the burrow entrance, a "latrine" sometimes occurs and often food
the hollows of riparian trees

remains are scattered here.

The temporary shelters occur under overhanging banks, in mill pond dams, under haystacks, etc. Daily activity and behavior. In the summer-autumn period, mink is active throughout the whole 24-hour period without a clear rhythm of daily activity. It more often hunts at dawn and at night. It is especially active in cloudy autumn weather with light rain. In winter, especially in severe frosts, it is less active. With availability of empty spaces under the snow, it does not come out from under the snow surface for many days. Activity of mink noticeably increases in autumn, when the young animals disperse, and sometimes, during the transition from completely frozen bodies of water to those conditions more favorable, and in spring during the breeding season.

The greater

part of the time,

mink go about on
rest,
its
it

the shores of

water bodies. Except for hours of

finds itself in continuous

are quick and

movement

in

search of food. All

it

movements

bustling. In case of danger,

tries to

hide itself in water, in extrees. It

treme cases,

it

conceals

itself

under roots or fallen

swims

and dives superbly. It was shown that it is able to run along the bottom of the water. While swimming, nearly half of its trunk appears above water. In hiding from enemies, it submerges under water leaving only the tip of its nose on the water surface. Diving, the mink is able to remain under water for 1-2 minutes, swimming in this time 10-20 m. After this, it appears on
the water surface for

poorly, but in pursuit,

It

2-3 seconds and dives again. The mink climbs it can climb a tree up to a height of four m.
it

hides well and observations of

rarely present themselves.

is

The mink
oped.

is

an animal in which sedentariness

well devel-

Due

to this, within separate bodies of water,

It

it is

very easily

killed out.

slowly resettles emptied places.

Seasonal migrations and transgressions. Lengthy migrations by mink have not been noted. In autumn, movements of separate

1104
individual are observed in connection with the changes in food

resources of areas and the freezing to the bottom of small forest

water bodies. In connection with the dispersion of the young, limited

wandering

is

observed. During these, mink pass from one

river or rivulet to another, traversing forest watersheds

which ex-

tend for several kilometers.

In spring,

male mink perform a quite long travels

district (P.B.

in search of

females. At that time, paths of minks are observed along the banks

of rivers of
735

Moscow

Yurgenson).

Reproduction. In
ApriH'.

Moscow

Zoo, estrus was observed on 22-26

Mating lasts from 15 minutes to 1 hour and proceeds several times in one day. Duration of pregnancy has been determined as 42-46 days. Parturition was recorded on 6 June. The number of young in the litter ranges from 3 to 7. Growth, development and molt. Weight of the newborn European mink equals 6.5 gm. The young grow rapidly, and after 10 days birth weight has trebled. Milk incisors appear quickly. They are born blind and the eyes open on the 30th-31st day. The lactation period lasts 2-2.5 months'*^, but at the age of 2025 days, the young already begin to taste food brought by the mother. Coming out of the burrow is noted from 4 to 27 July. On the hunt, they go out with the mother at the age of 56-70 days, and at the age of 70-84 days, they become independent. They attain half the size of adult animals by the end of July, but separate individual
of late
litters,

the size of a rat in August, occur.

Mink

attain sexual

maturity in the following year. Duration of

life is

undetermined.

As in other aquatic mammals, molt in mink proceeds slowly and gradually, and therefore is unnoticeable. Enemies, diseases, parasites, mortality, competitors, and population dynamics. The otter is the most dangerous enemy and competitor of mink. This is greatly weakened by the known degree to which the otter avoids littered waters and those overgrown with
aquatic plants.

The

otter

crowds out the weaker mink from

its

own

home

range. Those places where the

approximately one month
et al.,

number of

otters increases the

*H.e.,

(Rubetskaya

1933) that

this is

later than in the American mink. It is supposed one of the reasons for lack of success in attempts

at interspecific hybridization.

^Milk composition: fat

3.8%,

protein

6.2%,

sugar

5.6%,

mineral salts

10.66%.


1105

number of mink is greatly reduced. A series of cases of direct destruction of mink by otter are known. Inimical competitive interrelationships exist between both mink species. In areas where they live together, the larger and stronger American mink replaces
and often destroys the European mink. Competition also occurs with the forest polecat in those cases where it lives in the fioodlands of creeks. A case is known of a polecat set upon a mink and
dragged
736

burrow (Formozov, 1923). Diseases of minks in nature have not been studied. Invasiveness with helminth worms was determined as 56% (M.P. Lyubimov). Among helminths, 27 species were recorded in minks: trematodes 11 (Petrov, 1941). Particularly 14, cestodes 2, and nematodes often, pulmonary filariasis and krenzomatiasis are encountered, as well as skryabingulosis infecting the frontal sinuses. They someit

to its

times die in fishing tackle.

735

Fig- 262. Paired prints of

paws and schematic of mink jumping on snow. The

legs are

usually placed closely in series without skids, in contrast to ferrets and marbled

polecat in which the pair of limbs of one side are for ahead.
region,

Kostromsk

district.

Neya river, Ponazyrevsk 22 October 1939. Sketch by A.N. Formozov, about 2/3
natural size.

1106

Mink populations independently

harvested do not remain
in its variation has not

of the influence of those

at one level. However, a marked sequence been established. Usually, populations grow for 3-5 years. Such widespread death of mink is scarcely connected only with changes in abundance of food. Probably, climatic peculiarities determining the ice regime of water bodies are of

greater significance. In the severe winter of 1939/40,

when

all

bodies of water in Karelia were strongly frozen, the number of open-water areas was greatly decreased at the same time that mouselike rodents

were extremely few

resulting

in the majority of

mink

dying from hunger (Sludskii, 1953).

Field characteristics. Tracks of mink are nearly as large as
those of the forest polecat, but prints of the pads are larger and

more rounded, and the claws are shorter. Distance between the paws are wider than in the polecat, and track in the snow often appear dirty. The gait itself of the mink bears the stamp of restlessness and incessant roving from side to side. In the polecat, the gait is more stable and firm. In winter, the tracks of mink often sink deeply into the snow or are concentrated around open water and shoal areas. In rare cases only does it move away from water for more than 50-100 m (P.Yu.).
Practical Significance

The European mink is a valuable fur animal. During the period of most intensive trapping, the catch in the USSR reaches 50-75 thousand skins. At the present time, catch is regulated by issued
licenses.

The most

effective

means of capture

is

that utilizing

dogs (laika

or other breeds).

Mink

are also successfully caught with

jaw

traps

and box trap types, cage and plashka* and by obmet nets. During the hunting season, the experienced hunter catches 30-40 and up to 100 individuals. Spring harvest with the help of dogs is very

damaging
ing
is

in that period

when

the rising water

fills all

the

empty

places under the ice and drives the

mink

to dry land.

Spring hunt-

not permitted (P.Yu.).

*LocaI word for trap/lnet type

Sci. Ed.

1107

Subgenus of Polecats Subgenus Putorius Cuvier*, 1817

FOREST, OR BLACK, POLECAT^^
Mustela (Putorius) putorius Linnaeus, 1758
1758. Mustela putorius Linnaeus. Syst. Nat. Ed. X,
1, p.

46. Upsala,

Sweden. 5S. Mustela furo. Linnaeus. Syst. Nat. Ed. X, 1, p. 46. Africa, Morocco. 1827. Putorius vulgaris. Griffith, Cuvier's Animal Kingd., 5, p.

120. Substitute for putorius Linnaeus.

1843. Putorius foetidus Gray. List Spec.

stitute for

Mamm.

Brit.

Mus. Sub2, p.

putorius Linnaeus.
In:

1851. Putorius verus. Brandt.

Semashko. Russian Fauna,

357. Substitute for Mustela putorius Linnaeus.

1926. Putorius putorius stantschinskii. Melander. Nauch. izvest.

Smolensk, gos. univers., p. 137. Roslavl'sk gov. Described as a color "variety".

737
ta.

.,

Smolensk

1929. Putorius putorius orientalis. Brauner. Ukrainsk. mislivets

Ribalka, No. 2-3,
p. 9,

Ukraina. Exact type locality not

indicated, but apparently, northern part of former

gov.

Nom.

praeocc.

Khersonsk Mustela erminea orientalis Ognev,

1928.
1932. Mustela putorius rothschildi. Pocock. Scot. Nat. Edinb.

Dobrudza River, Romania.

1948. Putorius putorius L.
f.

piriformis. Kostron. Prace Mor.-slez.

Akad.

prir. ved., 20, p. 52.

Czechoslovakia.

1952. Putorius putorius ognevi. Kratochvil. Shorn. Vysoke skoly

zemed. a lesnecke v Brne, No. 1, p. 8. Middle European Mustela erminea part of USSR. Nomen praeoccupatum
ognevi Jurgenson.
1955. Putorius putorius orientalis. Polushina. Ekologiya, rasprostr.,
i

nar.-khozyaistvenn. znachenie sem. kun'ikh zap.-oblastei

p. 6.

Ukrainsk SSR,

Eastward of

line

Pskov-Minsk-Zhitomir-

*In Russian original, misspelled Guvier

Sci. Ed.

""Also

known

as

common

polecat (in books).

By

furriers

black

polecat.


1108
Vinnitsa.
orientalis

Nomen praeoccupatum Mustela erminea Ognev, 1928; Putorius putorius orientalis Brauner,

1929. 1965. Mustela (Putorius) putorius mosquensis. Heptner. Zool. Anz.,

176, H.l, p. 2. Savvino 20

km

east of

Moscow

(V.H.).

Diagnosis
General color brownish-black with light yellowish underfur visible.

Lower

side of

body

entirely blackish, without light under-

fur; large

yellow field absent on belly. Tail black throughout whole

its

length. Postorbital constriction weak,

width in narrowest place

not less or hardly less than interorbital constriction (V.H.).

Description

The general appearance,

arching
its

habits, etc. of the black polecat are typi-

cal of other species of the genus.

It also moves mainly in leaps, back high upward, or, slinking down, it creeps. However, it generally has a more compact conformation and, although short-legged, its body is not so elongated as compared even to the mink and steppe polecat. This partially depends, apparently, on strong elongation of the projecting axis, mainly in the posterior

part of the back. The tail is relatively short (about one-third of body length), covered its whole length by hairs of approximately the same length and slightly fluffy. Winter fur quite long, fluffy and soft, although not particularly close-fitting. Guard hairs along back elastic and coarse. Guard

on rump (in the sacral region); anteriorly, they become shorter, and shortest on withers. Fur on belly short, close-fitting. Average number of hairs per 1 cm^ on back 8,5009,000; ratio, one guard hair to 19-20 underhairs, and on abdomen
hairs particularly long

hairs

1 cm^ about 6,000. Length of guide on back about 50 mm, and thickness about 117 mk*; guard hair, correspondingly, 43 mm and 112 mk; and underhairs 26 and 20 mk (Tserevitinov, 1958). Winter fur of polecat is characterized by very great difference between length of guard hairs and of underhairs, which is not characteristic for the majority of species in the genus (except steppe polecat), and this causes guard

average number of hairs per

mm

*Microns.

1109
hairs to appear to protrude

above underhairs. This

tail at

is

further accen-

tuated by the contrast in color of the dark guard hairs and the light

underhairs (see below). The

base

is

covered by guard hairs

about 35

mm in

length and with underhairs of about 18

mm.

Else-

where on the tail, guard hairs are about 45 mm. Anal glands, giving a sharp "polecat" odor, are well developed (polecat may even project their secretion) and serve as a means of defence. There are 3 to 5 pairs of teats, more often 4.
738

General color tone of adult polecat in winter fur

sive brownish-black or blackish-brown,

is

quite inten-

which is determined by color of long guard hairs. Equally with this on dorsum and sides of body, the dark tone is brightened by bright whitish-yellowish, sometimes yellowish-grayish underfur showing through. What is perceived as bicoloration is obtained by what seems to be a twolayered color of the fur. The light undercoat
different parts of the body.
half,
is

not seen equally in

On

the back, especially the posterior

long and numerous guard hairs completely or almost comis

is

pletely hide the underhairs, and the light tone here

not noticewell defined

able or lightening

is slight.

On

the sides,

it

[lightening]

and

their general color is strongly differentiated

from the general

in color.

tone of the [dark] spine. Approximately the same relation occurs

on the neck. The nape and the shoulder region are darker

Fig.

263.

Black, or forest polecat, Mustela (Putorius) putorius L. Sketch by A.N. Komarov.

1110
Throat, lower neck, chest, belly and inguinal region are devoid

of light underfur
extent.

black or blackish-brown

throughout the entire

Sometimes, large blackish-brown chest and inguinal regions and the belly

on the is distinguished by its somewhat lighter, light-brownish tone. The chest and inguinal spots in this case are united along the mid-line of the belly as a narrow
fields are situated

black band. Ventral coloration characteristic of steppe polecat is, however, never observed (see [below]). Limbs are pure black or
with brownish
its

tint,

the

tail is

black or brownish-black throughout

is

whole length. Light underfur

absent on the limbs and

tail.

contrasting pattern occurs on the head

eyes, region between the eyes (anterior part of forehead) and longitudinal stripe along the top of the nose

the area around the are black-brown the

upper half of the forehead and the whole region between the eyes and ears, cheeks, around the mouth and chin are whitish or white, with a silvery tint on the forehead. Therefore, on the light head there is a "mask" covering the region of the eyes and the area

between them. The ears are dark-brown edged with white. Vibrissae are black. Hairs covering the lower side of feet are brownishblack or dark-brown; the digital and footpad callosities are
completely covered by them.

The general tone of

the fur

is

variables, occurring

from darker

to lighter. This mainly depends on the density of the guard hairs which cover, to a greater or lesser degree, the light undercoat and

on the intensity of

their black tone. Apparently, color

is,

to a

certain degree, associated with age

the older the animal, the red-

739

der the color tone of guard hairs and the lighter the general tone of the fur. The color of adult females shows no essential
differences, but, apparently, their underfur does not carry rusty

highlights and

on the whole whiter and paler. The summer fur is sparse and coarse, shorter than winter fur (guard hairs not longer than 30-35 mm), grayer, dull and lacking the beautiful luster characteristic of winter fur. Underfur is more weakly developed both in length and density and has a brownishis

gray or rusty-gray color.

Newborns are almost bare, but in the first days they become covered with short, fine and delicate, but dense, white hairs and they look whitish or almost white. This, apparently, is the delayed development of embryonic pelage (lanugo). At the age of 10 days,
this

pelage begins to be replaced by the second

the juvenile.

Its

1111
is short and sparse, but fluffy and soft, quite even over the whole body; guard hairs are not distinguishable ("down-like"). General tone of fur is grayish-tawny-brown with grayish-rusty underfur showing weakly on the sides. Lips and chin are white, but in some individuals, the head the facial pattern is very variable is uniformly dark, the same color as the trunk, and in others, a quite wide whitish band is located across the head between the eyes and ears. Animals found in the nest are in this fur, and have

fur

it

in the first

days after leaving the nest.

is

This pelage

converted in the
is still

first

summer

into the pelage of

subadult animals. The fur

somewhat "down-like", but guard

is "well

hairs are well differentiated

from underfur, the head pattern

defined, and contrast between color of underfur and that of guard

hairs
great.
is
It

clear. In general variation in color in this pelage is very

tially differ

first winter pelage which does not essenfrom the corresponding adult pelage (Herter, 1959,* Kratochvil, 1962 with modifications).

passes into the

Among

exotypic variations of polecat are

binos and in some places, the chromic deviation

schinskii,"
is

known complete known

al-

as

"braginskii polecat" to furriers, and described as "P. p. stant-

not

uncommon. These chromists vary

is

quite greatly in

color details. In typical individuals, underfur

brighter than in normal polecats.

usually reddish,

Guard

hairs

on the whole trunk

are bright and lustrous, reddish or brownish-red, or intensely red.

The
is

tail

is

reddish or reddish-brown. Black guard hairs are not

everywhere, being absent on the lower body and head. The venter
light,

having the undercoat color and only a large chest spot;

inguinal region and posterior part of

In

abdomen

are reddish-brown.

extreme cases, guard hairs are so

light that in tone, they are only

slightly differentiated

from the pale-yellow underfur or almost

is

indistinguishable. In this case, the whole animal

very light pale-

golden-yellow color, with only a small darkening on the chest

where

a large spot of light-tawny color is

[normally] located.

Equally with the very bright reddish polecats, animals are found
with normally colored underfur and only with reddish guard hairs.

They themselves represent

individuals and chromists.

a sort of transition

between normal

Geographic variation

in color is

weak.

Not

in Lit.

Cit Sci.

Ed.

1112

740

Fig. 264. Skull of forest, or black, polecat,

Mustela (Putorius) putorius L.

Skull relatively coarse and massive

more massive and heavier

and
in adult

than in mink.

It

is

comparatively short and broad, with a strong

but short, broad facial portion. Skull width in region of mastoid

processes more than half condylobasal length. Protuberances and

crests well developed; occipital crest large,

and old

1113
animals, sagittal crest well defined throughout whole length

as

well as anterior branches. Mastoid processes well developed.

Zygomatic arches strong, but not widely separated

zygomatic

width approximately equal to mastoid or only slightly exceeds it. Region between supraorbital processes is convex. Line of upper skull profile in facial portion descends anteriorly as a convex
arch, and the line of the braincase of skull gradually and slightly descends posteriorly. On the whole, skull somewhat flattened. Supraorbital processes small and protrude slightly laterally. Interzygomatic region of braincase relatively short and broad. Lateral outlines of postorbital portion form almost parallel or only

741

very slightly diverging lines; narrowest place of postorbital portion

not in form of sharp constriction

its

width here equals to or only

posterior to line uniting

slightly less than interorbital width. Greatest constriction of postorbital region with

jaws closed usually

is

lies

apices of coronoid processes of mandibles."*^

Nasal opening
terior half; carotid

usually less than height. Auditory bullae

somewhat compressed laterally and width somewhat swollen in pos-

foramen on inner side of bulla lies nearer its anterior end than its posterior. Ends of pterygoid processes have hook-like form and are strongly turned outward. Nasal bones widened anteriorly and posteriorly constricted, forming acute wedge passing between anterior portions of frontal bones. Therefore, upper
part of premaxilla

comes

into close contact with nasal, but not

latter (noticeable

throughout whole length of

animals).

only in skulls of young

By comparison

with previously examined species in genus, teeth

very strong, large and massive in relation to general skull dimensions.

Camassial teeth relatively

large,

upper molars, on contrary, with smaller

all

dimensions. In large individuals,

described characteristics of skull

more

strongly developed and defined than in small; the latter reveal

certain arbitrary infantile features (see below).

Sexual dimorphism in the skull, not to speak of dimensions, is quite significant. Female skull lighter with less developed protuberances, crests, etc., not so wide and angular, and

more narrowly

separated zygomatic arches. Supraorbital processes usually smaller,

ferentiating

"According to Stroganov (1962), this is one of the most constant features difit from the skull of steppe polecat. Judging by material in Z[oological] M[useum of] M[oscow] U[niversity], it has no such significance.

1114
nasal openings narrower, convexity of skull in interorbital region
less, teeth

weaker.
(less than

one year), cranial region of skull convex and does not bear crests; supraorbital processes weakly developed, postorbital constriction weakly defined and narrowest place in this part of braincase lies almost directly behind supraorbital processes. Facial portion of skull and nasal region very short, auditory bullae relatively small; sometimes interparietal bone noted. All these features are more pronounced in younger animals. However, course of age changes in the polecat skull is quite great and comparatively prolonged. One-year-old animals greatly differ from older ones, and the two-year-old
In

young animals

relatively

much

larger,

animals are well distinguished.

Geographic differences

in skull are not manifested.

form typical of other species of the genus. Its base is somewhat compressed laterally, tip of bone is bent upward in form of a hook and somewhat twisted to the left. Ventrally, it bears a quite deep groove, continuing on upwardly bent tip. Left side of groove wall slightly elevated and forms a rounded protruding bony flange on very end of hook

The OS

penis, in general features, has a

(Ognev, 1931).

Caecum

not developed. Length of male intestine averages about

235 cm, of females about 186.5 cm, weight of male heart averages 7 gm, that of females 4.75 gm (Middle Europe; Herter, 1959).* Dimensions of the black polecat are subjected to very great variation, on the whole greater than in the previously described species of the genus. This is explained by the considerable difference in dimensions of males and females, and the comparatively
742

slow maturation of the young. They attain complete growth, apparently, only in the third year of life. Equally with this, is magnitude
of individual variation in both males, and also females
cially the former. Besides these

but espe-

"normal" fluctuations

in variation,

both relatively very small and very large individuals (three types according to measurements already mentioned by Hensel, 1881)
are

met with among

lie

polecats.

Measurements of the

latter ("giants")

are considerably larger than those of polecats of the "normal" type,

and

beyond the

limits of usual amplitude of their variation,

although. connected with them, representing rare positive variants.

*Not

in Lit.

Cit. Sci. Ed.

1115

The same applies

though
in the

to very small individuals.

The forms of these

of the genus,

three types are also

known

in several other species

one described here, they occur more often (see also

section on steppe polecat)/^ In

some cases

the possibility of hybrid

heterosis (with mink) cannot be excluded; however, this has not

115-167 mm, of females, 84-150 mm; length of hind foot of males is 42-63 mm, of females, 33-58 mm; ear length of males is 21-36 mm, of females, 15-26 mm.'*' Condylobasal length of skull of males is 59-71.2 mm (65.2), of females, 51-62.9 mm (55.7); zygomatic width of males is 35is

been remarked on for "giants." Body length of males is 350-460 (up females, 290-394 mm; tail length of males

to

480

?)

mm,

of

42
is

mm

(34.4); interorbital width of

males

is

15.8-20.3

mm

(17.5),

of females, 12.6-18.1

mm

(14.0); postorbital constriction of

males

14.3-18.0

mm

(16.1), of females, 12.6-15.8

mm

(14.3).

Length of adult os penis (8) is 33.5-M36.5-38.2 mm, of young 31.0-M34.0-37.0 mm; weight of penis bone is 0.280-M0.3370.470 gm, of young, 0.080-MO. 146-0.240 gm (Popov, 1949). There are no reliable data on weight of our polecats (clear errors are sometimes found in literature, for example Stroganov, 1962). Weight of adult males from Middle Europe are 1,000-1,500 gm, females, 650-815 gm (Herter, 1959) (V.H.).
(17),

Systematic Position

The steppe

polecat,

M.

(P.)

black polecat. Their closeness

eversmanni is the closest form to the is undoubted and is generally so

great that a series of authors, beginning with

Pocock (1936)
and

in-

clude them in one species. However, there

principle for this and, evaluating
sides of their existence,
it is

is

insufficient basis or
all

all

their particularities

more

correct to consider both polecats

as a closely related, but independent, species (for details, with

'The above described extremely wide variation in weasels of the European part

of the

itself represents a phenomenon whose nature is of a different kind. Ognev (1931), Heptner and others (1950), Polushina (1956) and Stroganov (1962) with some additions and corrections, from materials in Z[oological] M[useum of] M[oscow] U[niversity] and others. With skull measurements, average size of Middle

USSR

"'After

Russian polecats after unpublished data of L.G. Morozova-Turova (40 adult males, 24 females) are given in parentheses.

1116
analysis, see systematic position of white* polecat below).
this, the

From

black polecat must, apparently, be considered a less spe-

cialized

form with several features of an

infantile character. In

skull structure, adaptation to a predatory

mode

of

life, in

particular

to capture of larger animals, did not reach the high level of the

steppe polecat.

The

relationship between the black polecat and

ciently close, although at a lesser level.

represents, in a certain sense, a

mink is suffiThe European mink itself connecting link between polecats

and the remaining species of the genus (for details see the sections on "Systematic Position" of European and American minks). This offers no difficulty, as already mentioned, except for the rarity of
hybrids of both species in nature. In recent times, the suggestion

was even made (Stroganov, 1962)

743

to include

minks

in the

genus of

polecats (Putorius) as a separate subgenus.

In

West and Central Europe,

a kind of domestic animal for the

is

destruction of rats in houses and for hunting rabbits

is

kept,
is

which

called the white polecat, Mustela furo.** This

its

form

a typical

polecat in

general appearance but

is

pure white in color with

red eyes (albino). The origin of, and systematic relationship be-

tween the white polecat and the European as well as black and steppe polecats has been studied frequently, but this question has not been resolved. Some consider it a domesticated albino form of

M. putorius and others

of M.

eversmanni.

It

is

also suggested

that the white polecat is a special

North African form of the black polecat which has been preserved only in a domestic condition. In the majority of cases, this form is recognized simply as "M. furo"
or "P. furo'' (for details, see Herter, 1959). White polecats are,

apparently, unrestrictedly fertile in crosses with both the forest and

steppe polecats.

The
cat;

skull of

"M. furo"

is

generally typical of the forest poleit

however,

in several cases,

bears features characteristic of

It

the steppe (constriction of postorbital region).

that hybrids of black polecats

has been noted

and "M. furo" closely resemble the Siberian polecat in color (color of lower body). Apparently, it is more correct to consider that "M. furo" is a
domestic form of M. putorius. This
is

indicated not only by

to

its

Alternate name for steppe polecat, but also applied

below) Sci. Ed. **ln English, "ferret"

domestic ferret (see

Sci. Ed.

1117

morphology.

".

furo" appeared

in

southwestern Europe (appar-

ently on the Pyrenean Peninsula) where only M. putorius lives and

has lived, and, apparently, from a form which lives (lived?) in

extreme northwestern Africa {M.

long ago, in a

p.

furo

L.,

Western

Rif; accord-

ing to Cabrera, 1932; Allen, 1939), This, apparently took place

much earlier epoch, when it hardly seemed possible M. eversmanni could be brought so great a distance to be domesticated. Moreover, there is also no basis to speak of M. furo as an independent species. Some similar features between the skull of M. furo and M. eversmanni are natural, a result of the close relation between M. putorius and M. eversmanni. This may also be the result of some deviations in the skull structure of M. furo as a result of prolonged domestication. Its hybridization with M. eversmanni is the same as the hybridization with M. putorius, but this possibility was extended by domestication (V.H.).
that

Geographic Distribution
Forest, forest- steppe, and in part steppe zones of the major parts

of Europe, and extreme northwestern Africa.

Geographic Range
This

in the Soviet

Union
of the species range.

itself represents the eastern half

The western border of

the range of the polecat in our country

constitutes the state frontier,

from the mouth of the Donau [Danube] in the south approximately to 6240' N. lat. (northwest of
Suoyarva), on the border with Finland in the north.
In Karelia, the northern border extends

scribed above

mately to the
the

from this point desomewhat towards the southeast at first, approxiSpassk Bay of Lake Onega, thereby passing around
the south and then, passing around
it

West Karelian uplands from

these uplands from the east,

suddenly ascends directly to the

lat.

north passing in particular, near the western shore of Segozer and

reaches Rugozer

at 64*'10'

N.

Thence, the border line turns

somewhat northeast and crosses the Lekhta, reaching Kem' on the White Sea (65 N. lat.). This is one of the most northerly points of species' occurrence. The data given apply, on one hand, to the early 1950's, and on the other, concerning the Rugozer- Kem',

1118
to the

end of
is

it

(Marvin, 1959; Parovshchikov, 1959). Since the

it

polecat

spreading northward,

is

possible that

its

distribution
sixties

in eastern Karelia after the fifties

and the beginning of the

is

has changed and the border extends farther north than

shown.

The White Sea coast constitutes the northern limit of the range throughout its whole extent from the mouth of the Onega to the mouth of the Severnaya Dvina. From Arkhangel'sk, the border goes to the city of Mezen' at the mouth of the Mezen'. In the expanse
744

between Arkhangel'sk and Mezen', the range, apparently, reaches the

at the Arctic Circle attains the other most by the species (Mezen', about 65 N. lat.). From the mouth of the Mezen', the border suddenly returns to the south and approaches closer to the upper Mezen' in an unclear way (probably along the Mezen') at a place near 64' N. lat. Thence, the northern border of the range goes on, to the upper Vychegda (near 63 N. lat.; V.Ya. Parovshchikov). Farther, the border apparently descends steeply to the south and in the Urals, lies at 58 N. lat. or somewhat more southerly (Shvarts, Pavlinin and Danilov, 1951). However, judging by its distribution on the Vychegda, the border may actually be more northerly.

seacoast and here

almost

northerly point inhabited

745

The eastern border

is

very poorly known. Apparently,

its

it

ex-

tends along the Urals, perhaps even to

eastern slopes, embracits

ing Sverdlovsk from the west. Old information on

the Irbit region require confirmation.

occurrence in

According

to recent data, "forest polecat is absent in Trans-

Urals (to the east of Sverdlovsk, V.H.) at the present time"

(Stroganov, 1962). Indications concerning former Tyumen, Ishim,

Kurgan and

regions (Slovtsov, 1892) clearly apply to the

steppe polecat, although these are given for "Foetorius foetorius".

The

distribution of polecat southward along the Urals

is

un-

clear. In the southern Urals (south of the latitude of

this species, apparently,

Magnitogorsk),

does not exist and here only the white found everywhere (Kirikov, 1952). Information concerning the occurrence of the described species along the eastern slope
polecat
is

of the Urals nearly to Orsk (Bobrinskii, 1944)

is

incorrect.

The southern border of the range begins in the west at the mouth of the Donau [Danube], extends eastward along the Black Sea coast to the mouth of the Dnepr, whence it moves back from
the Black Sea coast (to the latitude of Askaniya-Nova) reaching

1119

1^].

1^3

744

Fig.

265. Borders of the ranges of polecats in the European part of the

USSR.

1 range border of forest polecat, Mustela (Putorius) putorius L., 2 western range border of steppe polecat, M. (P.) eversmanni Lesson, outlining the area of sympatry of both species, 3 place of occurrence of forest polecat at Achikulak in the

V.G. Heptner.

Cis-Caucasus. Question marks in the Trans-Urals, Irbit, in Kareliya confirmed settlement in 50's and beginning of 60's.

area

of

1120
the shore of the

Azov

Sea, and along

is

it,

goes to the mouth of the

Don. The black polecat

polecat
is

absent in the Crimea.

In the steppes of the

European part of the USSR, the black rare everywhere, and has a very unique and limited

distribution

it

is

associated only with forest tracts, in part with

human

settlements and with the edges and valleys of rivers.

The
pole-

farther to the south, the stronger this association.

cat lives in the

The steppe

open steppe, and

in

forest very rarely occurs

(Askaniya-Nova).

mouth and lower Don, the range passes into the The southern border in this section passes along a line from Stavropol to Krasnodar the western border being formed by the Azov Sea, and the eastern, a slightly curved line directed from Stavropol northwards to Tsimlyansk district on the Don'*^. This latter quite closely
the

From

steppe of the western and middle Cis-Caucasus.

coincides with the borders of the Cis-Caucasian steppes.

Details of the border of the range between the
are not fully clear. Apparently,
it

Don and Volga

suddenly begins to ascend northto the

ward along the Don, and then passes

of Saratov.

Volga somewhat south

In the Saratov steppes of the Trans-Volga, the polecat is

absent

it

is

encountered only

in the

extreme lower B[olshaya]

steeply returns

and M[elenkaya]
the Volga.

Irgiz. Farther, the

border goes to the north along

it

Somewhat

south of the Samara bend,

to the east and, passing

53**

around Obshchii Syrt approximately along

at the latitude

N.

lat.,

reaches the Urals

of Magnitogorsk.'*^

occurrence on the
data

"The Cis-Caucasian part of the range is outlined here according to the individual map of Vereshchagin (1956). These points, in turn, were based on of fur manufacturers. It is not excluded that in some cases, skins were brought

from the north. In any event, the black polecat is extremely rare everywhere in the Cis-Caucasus and individuals are met with among masses of steppe polecats. Earlier
(Satunin, 1915),
its

presence in the Caucasus was denied and

is

is

sometimes

still

denied

(Kuznetsov, 1952). The possibility

individual animals

not excluded that this species

may

penetrate as

somewhat farther to the southeast; for example, it was recorded in Achikulak about 240 km to the east and southeast of Stavropol (V.G. Heptner). "'Range after data of Bogdanov, 1871; Slovtsov, 1892; Brauner, 1912, 1914, 1928, 1929; Lobachev, 1930; Ognev, 1931; Isakov, 1939; Bobrinskii, 1944; Vereshchagin, 1947, 1959; Pleskii, 1941; Kuznetsov, 1948; Heptner et al., 1950, 1956; Elpat'evskii, Larina and Golikova, 1950; Marvin, 1951, 1959; Shvarts, Pavlinin and Danilov, 1951; Kirikov, 1952; Komeev, 1952; Sludskii, 1953; Novikov, 1956; I. P. Laptev, 1958; Parovshchikov, 1959; Afanas'ev, 1960; Sokur, 1960; Stroganov, 1962 and others and according to materials of M.Ya. Marvin, V.Ya. Parovshchikov, N.I. Larina, L.G.Turova and V.G. Heptner.

1121

The range of black polecat

746 mately),
i.e.

to the south

of the line Zhitomir-

Kiev-Orel-Voronezh-Gor'kii-Kirov-Ufa-southern Urals (approxiin its greatest part,

overlaps the range of, the steppe

polecat. In this region, individual hybrids are encountered (quite

rarely), but hybrid populations are not

The black polecat

ably widening
its

itself represents a species

Its

formed anywhere. which is notice-

range towards the north.

movement

is

ob-

served in Finland (Kalela, 1952), in Karelia (Isakov, 1939), Arkhangel'sk district and Komi ASSR (Lavrov, 1925,
Parovshchikov, 1959), and Kirov district (Lobachev, 1930). There was information that it also colonized eastward to the Urals and

beyond the Urals, but

it

was decided

that this animal passed be-

Fig. 266. Colonization of the black polecat to the north in the Karelian

ASSR

(Isakov,

1929; Marvin, 1959) and in Finland (Kalela, 1952) with additions. V.G. Heptner.

1122

yond

the Urals not long ago.

its

However, there are no accurate data

to the north
is

concerning

movement

in this direction.

The tempo of colonization

towards the northwest about 250

quite energetic.

Thus, in Karelia, from 1930-1932 to 1956, the polecat colonized

km

(from Vodlozer to Rugozer).

In Finland, in recent decades the polecat settled nearly all over the

southern half of the country (Kalela, 1952).

From 1930-1932

to

1956, the polecat settled along the

latitude of

Severnaya Dvina from the

i.e.

Shenkursk

to

Arkhangelsk,

for a distance of about

320

km

in
at

a straight line (Parovshchikov,

appeared

Kaisk Pochinki on the

Kama

at

1959). The polecat 60 N. lat. at the end

it

of the 20's of this century (Lobachev, 1930) and by 1956,

began

to

be found, though not often,

it

in

the upper

Vychegda
north-

(Parovshchikov, 1959). Therefore,

years. Within 25 years,

moved about 300 km

the Onega.

ward or northeastward during a period of approximately 22-25

it

a
is

tempo of colonization
Colonization

on average about

moved 300 km along

Such

10-12

km

per year

considered to be very high.

is mainly stimulated, apparently, by felling of forand ploughing. However, in some places, it occurs in regions where there are no essential changes taking place in the landscape. Apparently, a certain role is also played by climatic warming which has occurred in northern Europe in places in the last 100 to 150 years. It is considered to be the main cause in Finland. Besides colonization of new places and progressive movement of the bor-

ests

der in several segments, a more or less significant "oscillations" of

the border occur locally.

Geographic Range outside the Soviet Union

This occupies almost
of Europe. In the north, the border in66 cludes Finland south of N. lat.; in the Scandinavian Peninsula, it begins at the Baltic coast at almost 60 N. lat. descends a little
all

westwards including Lake Vattern* from the south, in southeastNorway, it forms a narrow projection towards the north approximately to 61 N. lat. In the west, the range includes England (absent in Ireland) and in the southwest, it includes the Pyrenean Peninsula and the Rif region in Morocco. The southern border passes along the coast of the Mediterranean Sea (absent on the
ern
*In Russian original, "Venern"

Sci. Ed.

1123

^
the Dobruja region (V.H.).

Balearic Islands, Corsica, Sardinia and Sicily). In the Balkans,

the range includes the northern part of Yugoslavia; eastward, the

border passes along the Donau [Danube], encompassing, however,

~4^

1=

=4

jA

^'^

J^

Xj^-

^.
=s^

^^^'^^^
'''
'^

b^
^
^""^"^ Mezen'
I

^^^^
V-.7-

"""--^^V,^^^^_^^_^ '^
-:

[^^^

^^

(:
-f'""

:-

/^

^ :

/^
\

1944
)

^\

TTI^

7-^

l/^^k

"^T",--^: ,/^^-='-k

^^
^
/\

^^Unskay^M^^^'^'lH

959

^^v

iL,""^^

hi^^^*"^

"V

v^~\.

/^

r^

747

Fig.

267.

Movement of
district

the northern border of the range of the black polecat in

(Parovshchikov, 1959, with additions). Lines designate the general limits of the range-points the individual appearance of animals far from the

Arkhangel'sk

general border.

The movement of the border does not fully correspond was established for Karelia. V.G. Heptner.

to that

which

1124

748
\

Fig.

268.

Species range of the black polecat, Mustela {Putorius) putorius L. V.G. Heptner.

Geographic Variation
Features of the black polecat geographically change negligibly
within the borders of our country.
polecats of the steppe zone are
forest
It

is

possible that the black

somewhat lighter than those of the zones (Brauner, 1929). However, these differences apply

only to a very small part of the species population in the

southern population does not deserve special designation.

USSR;
If this

they are not yet proved sufficiently and are so insignificant that the

form

is

actually distinguished,

it

is,

apparently, identical with the

1125

race rothschildi described from Dobruja (the

name

orientalis, of

Brauner, though earlier was preoccupied, see above in synonymy).

The polecats of the middle and eastern districts of the European part of the country differ somewhat from the western and middle European ones, as well as those occupying the most
western parts of the range with us. This, in one form or another

was

stated

much

earlier (Satunin,

1895; Brauner, 1929; Ognev,

1931), and in recent times, this was confirmed by

some

authors,

who

formally distinguished the East European polecat (Kratochvil,

1952; Polushina, 1955).

series of unclear points

were associated

with the nomenclature of this form, and thus there was a need for
749 a

new name,

since the suggested names, including aureus, do not

apply to the middle Russian polecats (see note in synonymy of

steppe polecat, page 1134 footnote 51 and also Heptner, 1964, 1965).

Within the boundaries of the range of polecats

part of the

in the

European form are

region),

USSR, some

geographical localization of the red muparticular, Roslavl'

tant "P. stantschinskir is recognized. Individuals of this

known from Smolensk district (in Tambovsk, Moscow district (Klin

below Ostashkov, Pereslavl'-Zalessk (coll. Z[oological] M[useum of] M[oscow] U[niversity]) and Byelorussia. The name itself, "braginsk polecat, given to this form by fur manufacturers for the city of Bragin
region),

(southern Byelorussia), points to a certain geographical restriction.

Within the USSR, two races of the black polecat may be

recognized.
1.

Middle Russian black polecat, M.

(P.). p.

mosquensis Hept-

ner,

1965 (syn.

orientalis Brauner, orientalis Poluschina, stants-

chinskii, ognevi).

Body and

skull

dimensions relatively small, fur relatively

light,

slightly fluffy with insignificant luster.

Found in European part of USSR east of line passing approximately through Pskov-Minsk-Zhitomir- Vinnitsa.
Outside the

USSR absent.

Source of above description given of the species is mainly according to material of this form; measurements are of Moscow
polecats (see p. 1115),
2.

Western black polecat. M.

vulgaris, foetidus).
skull

(P.) p. putorius Linnaeus,

1758

(syn.

Body and

dimensions larger than Middle Russian form,

fur darker, lustrous and fluffy.


1126

Found west from

in

western part of European territory of

i.e.

USSR

to the

the given line,

Pribaltika;

Western Byelorussia,

western Ukraine.

and western Europe. Differences between the above-mentioned two forms are real, but they are not sharp and are mainly revealed in examination of a series of them. It is entirely probable that characteristics of Middle
Outside the
in central

USSR, found

Russian polecat are associated with its relatively recent colonization of the territory it now occupies (colonization beyond the Ural
is,

apparently, continuing).

Outside the
1)

USSR,

the following forms are usually recognized:

M.

(P.) p.

furo Linnaeus, 1758

extreme

northwest of Africa

(apparently, does not presently exist in wild state; see above "Sys-

tematic position"); 2) M. (P.) p. aureola Barrett-Hamilton, 1904

Spain; 3) M. {P.) p. rothschildi Pocock, 1936

4) M.
(P.) p.

Dobruja, Romania;
5)

angliae Pocock,

1936 England;

M.

(P.) p.

caledoniae Pocock, 1936 Scotland. The differences between these forms

are, apparently,

very slight

and require further study. (V.H.).

Biology
Population. Concerning populations of the forest polecat and
distribution within the range, these
its

may be judged by

indirect data

the figures of

commercial catch. Before the October revolution, this animal were taken in Russia though locally it was not exploited. In the period up to the Great Patriotic war of 1941-1945, level of the catch increased about 2.5 times on average, and in some places, overexploitation was observed. A rough approximation of the number of polecats in the USSR might
about 150 thousand skins of

be estimated as 200-400 thousand animals.

The

distribution of population density within the range

may

be determined by indices of skin yields in 10 km^. Before the revolution, this yield from the entire range was 0.40, and in 1928/29 1.10. The greatest quantity of skins was obtained from

Byelorussia. Smolensk, former Valikoluksk, Leningrad,

750

Moscow,

and Ivanovsk

districts

and from the Ukraine. Indices of catch from

1127
these ranged (in 10 km^) from 0.028 in former Severnaya Territory to 4.18 in former Zapadnaya district, 6.28 in B[yelorussian] SSR, and even to 8.70 (region of Verei city, Moscow district). The latter figures already clearly indicate overhunting then. In Germany, these indices fluctuated between 0.43 and 0.77. The forest polecat there was strongly extirpated. Therefore, the greatest density within the

boundaries of the
is

USSR

is

observed

in the

western part of the range, which

Habitat.

explained by the favorable

habitat conditions for this animal there.

The

forest polecat avoids large, continuous forest

massifs. Small forest islands and isolated groves alternating with

meadows, cultivated
able for
its life. It is

fields

and human settlements are more

suit-

a typical representative of the belt of forest

islands and a true "fur-bearing" animal.

Within the boundaries of the taiga forests of the European north, a few polecats are encountered in squatter's settlements, and
along the banks of shallow

swamps and
it

forest creeks,

and they are

generally restricted to relatively populated regions. Together with

human
taiga.

cultivation and agriculture,

gradually penetrates into the

In the middle zone, typical habitats of the forest polecat

com-

prise flood lands of small rivers and creeks with their lakes and

swamps

the so-called "water"

meadows and

water-saturated,
it

boggy

depressions. Without penetrating deep into the forest,

adheres to

openings, borders scrub forest and old clear-cuts, especially those

adjacent to worked fields and settlements. Rarely
it

with in mossy bogs or in deep forests.

It

settles

may be met more often in

barns, storehouses, in cattle yards, in cellars of occupied houses,

villages and even the outskirts of cities (even cities as large as

Moscow).
In the forest-steppe

and steppe

belts, the forest polecat is re-

woody ravines and gorges, and the shrubby floodlands of rivers. Following these latter, it sometimes penetrates deeply into purely steppe regions. Food. The main food of the forest polecat, everywhere they occur, are mouse-like rodents. Among them, the gray (common)
stricted to settlements, isolated farms,

vole [Microtus arvalis] occupies

rence).

first place (18.5-26.8% occurRed-backed voles [Clethrionomys] are more rarely encoun-

tered (4.4-5.5%). In the floodlands of large rivers, the water vole

[Arvicola] acquires essential importance (average occurrence about

1128

15%). Of great significance in food of the forest polecat, espemainly, grass frog and green cially in winter, are amphibians

These foods have less caloric value, and therefore the polecat never becomes fat from them even when they are plentiful. Birds occupy third place in food (8-16% occurrence): domestic hens and pigeons, quail, gray partridge, grouse and various
toad.

small birds. The role of the forest polecat in the destruction of

poultry
in
is

greatly overestimated. Special investigations

showed

that

kolkhozes [cooperative farms] and poultry sovkhozes

it

[state

farms], losses from

ranges from 0.2 to 2.5% for the adult birds

pullets,

and from 0.5 to 12%, for

most often

in the

haying period.

Among
insects.

number of

rare foods of the polecat are the hedge-

hog, adder [Vipera aspis], grass snake [Tropidonotus natrix] and

The

significance of individual foods changes depending

in

on fluctuation

abundance of the main food

mouse-like rodents.

In case of their insufficiency, the significance of the less valuable

food (amphibians) or that of secondary importance, including carrion, increases.

In connection with

unevenness

in distribution of

mouse-like

rodents, the composition of food and their ratios are usually differ-

ent in the separate regions and districts (see Table 68).

In poorly studied

summer foods

in

flooded areas, the water

vole predominates (89.1%), further followed by small voles and

mice (28.8%), and

751

fish

(0.9%) (Grigor'ev and Teplov, 1939).

(in

Table 68. Geographic variation in foods of the forest polecat occurrence)

percentage of

Type of food

Volga-Kama

Territory (Girigor' ev
1

Middle zone
(Lavrov, 1935)

and Teplov, 1939)

Summer

Wintei
lim

Winter

M
Mouse-like rodents
28.8
89.1
squirrels

M
25.4
14.7

M
51.2
2.4 2.4
16.3 19.5
0.8 9.8

Water vole Hamsters and ground

Insectivores

Birds

Amphibians
Fish

Carrion

0.9

42.065.0 3.732.1 3.39.8

08.0 012.2 020.0 016.0

4.1

8.0
12.1

6.9

Number of

records

114

563

123

1129
In the western districts of the U[krainian]SSR (analysis of 11

stomachs and 31 feces), mouse-like rodents predominated in the food of the forest polecat, though the occurrence of all 18 components was not higher than

10%

(Polushina, 1958).

%
Hedgehog Shrew
1.8

%
Gray vole Domestic rabbit Sparrows Song birds
Bird eggs
7.6 7.6 7.6
5.0
3.4 9.3

2.6
6.8 7.7

Norway rat House mouse Harvest mouse

Red-backed vole
Mice, Apodemus

0.9
3.4

Amphibians
Fish

6.0
3.4
1.8

4.9 4.2
5.0

Water vole Root vole

Carrion
Insects

The

forest polecat characteristically

makes food

stores.

The

bodies of water where grass frogs overwinter are used as "store-

houses" by polecat
visits

in the winter time.

The polecat continuously

such places.
range. In contrast to the steppe polecat, the forest pole-

Home
cat lead a

more

settled

way of

life

and has a definite individual

home
nor
is

range. However, the dimensions of the range are

there data on length of
its

unknown

daily route. Polecats living in

villages, among buildings, have a very small home range. In February-March, as a result of food insufficiency, dimensions of the home range noticeably increase. A case is known when the

polecat

moved about

km

during a night (Lavrov, 1935).

Burrows and shelters. Forest polecats rarely dig burrows. The permanent burrow dug for them has a simple structure one short and shallow passage and a small nesting chamber. They sometimes

settle in

burrows of badger or fox. More often, the polecats use

natural hiding places as permanent or temporary shelters

heaps

of brushwood, wood-piles, old stumps, haystacks, etc. In villages,

the polecat settles under the floor of stables, saunas and other

inhabited and uninhabited buildings, and in cellars and storehouses.

Daily activity and behavior. The forest polecat

rarely active during daytime, only
is

is

an animal
It is

with a crepuscular and nocturnal rhythm of daily activity.

very

when motivated by hunger. This confirmed by laboratory experiments (Kalabukhov, 1943): the

daily

rhythm of the

forest polecat

was shown

to

be monophasic,

with the active period after 20:00 hrs with some reduction at midnight. The indices of the activity of young polecats (2.0-2.5
752

months) differ greatly: it is equally active throughout the course of 24 hours the day and night, with some increase during daylight hours. The activity of young females during daylight hours is less
expressed than in young males.

European polecat

is

bold and malicious*.

It

climbs, swims and

digs in the earth well, but rarely uses these abilities, being a
typical terrestrial carnivore. Its running
is

less

complex and

twist-

ing than that of

mink and ermine.

It is

also less quick and evasive

in condition

than solongi, ermine and weasel

man

can catch a
it

running polecat. The sense organs are well developed but

distinguish between different colors.

cannot

Seasonal migrations and transgressions. Only small seasonal

migrations are

tumn, many polecats

known but far from all move to villages,

polecats participate. In auattracted

by the concentrais

tion here of mouse-like rodents after the gathering in of the harvest

from the

fields. In spring (April), a reverse

movement

observed.

Reproduction. As
activity extends

in the steppe polecat, the period of sexual

from the second half of February

is

to the

second

half of June. Evidently, the rut often extends into

April-May

(Lavrov, 1935). Duration of pregnancy

Manteifel', 1947).

42 days (Moscow Zoo;

Body length of newborn animals is 7 cm, weight about 7 gm. Number of young in a litter ranges from 2 to 12, most often 4-6
(Lavrov, 1935).

Growth, development and molt. Young polecats are born blind

with the ear openings closed by a flap of skin, and covered by

down. Eyes open on the 34th-36th day; at wool darkens. Sometimes, the brood does not disperse until spring. Sexual maturity is attained at about one year, but full development and dimensions at two years. Molt in the in spring and autumn. forest polecat occurs twice annually
sparse, short whitish
that time, the

Enemies, diseases, parasites, mortality, and competitors. Competitors may be all carnivorous animals and birds feeding on mouse-like rodents: mink, ermine, fox and in recent time, raccoon

*Sic; Russian

word

is

zloba

Sci. Ed.

1131 dog;

among

birds of prey

the buzzard and owls and also viper

and other [snakes].

In the forest polecat, various infectious and parasitic diseases

from causes that are usually unmore often found than other carnivores. The forest polecat is susceptible to canine distemper, and it suffers from an unknown infectious intestinal disease accompanied by high mortality. A number of helminths were recorded, including not less
occur. Bodies of polecats, dying
clear, are

than four forms of nematodes parasitizing the frontal sinuses and

nasal cavity. In France, distomatosis

was revealed
fleas

in

every fourth

and causes and ixodid mites occur on the forest polecat. In captivity, the forest polecat lives up
animal. In
cases, the infection reaches the brain

some

the death of the animal. Usually,

many

to 12 years (ManteifeF, 1947).

Population dynamics. The forest polecat

unstudied.

is

characterized by

significant fluctuations in numbers. Their cause and character are

From 1928/29

to 1949/50,

i.e.

for

22 years, numbers of
in the

forest polecat in the southwestern part of Kalinin district increased

three times, but to different levels.

The increase
rise in

number

in

1945/46 was three times smaller than the

times smaller than the

1929/30 and six

maximum
is

rise of 1938/39. in the

There
lation

is

no

strict periodicity

population fluctuations.

Apparently, the period

about 7-9 years. Reduction in the popu-

may

last

from 3

to 5 years,

and the growth phase

in

both

mentioned cases equaled 3 years. The sharp and rapid population

reduction after 1938/39 occurred
at

a time of severe

summer drought

and hard winters of 1939/40 and 1940/41, which caused a sharp and deep depression in the mouse-like rodent population. As a
result, the

frequency of track occurrences of polecat

1940/41
fell 9-fold.

in the period

1939/40

to

During 22 years,

it

fluctuated from

0.03 in 10

km

in

1946/41 to 6.03 in 1938/39,

i.e.

by more than

200-fold. In the post-war years, the polecat population (as well as

of ermine) was low everywhere, but the cause of this was unclear.
753

Field characteristics. Tracks of the forest polecat are 1/3 smaller

than marten,

its

jump

is

equal to 40-65

callosities of the feet are very distinct

cm and prints of claws and (Formozov, 1952). For difWhen

walking,
it

ferences from tracks of mink, see page 1106. Triple prints of the
feet are as typical for the forest polecat as pairs.

frequently changes feet, and often one pace for another.

eral character of the track or trail is distinguished

The gen-

by an absence

1132

753

Fig. 269.
river,

Tracks of the hind and fore feet of black polecat on silt. Shore of Moscow Zvenigorod. 22 June 1939. Sketch by A.N. Formozov, about 2/3 natural size.

of elaboration typical of the mink.

in

It

moves

in energetic leaps, digs

places inhabited by mouse-like rodents, and prefers to catch

in the

animals moving on the surface, so rarely digs

is

snow.

Its

track

not so tangled as weasel and ermine (Formozov, 1952) (P.Yu.),

Practical Significance

The

forest polecat
is

is

a valuable fur-bearing animal.

The

price of

its

skins

higher than that of steppe polecat. The fur enjoys a good

the world fur market on which, before the

demand on

1914-1918
October
increased

war, Russia offered more than

50%

of

all

skins. After the

revolution, the take of this species in the whole

USSR

noticeably as reckoned by growth of the catch, and also as calcu-

number in western European countries. Second World War, our population of this species decreased somewhat. This was reflected in the volume of catch of the forest polecat. In 1956-1969, as compared to 19261929, its take decreased on average in the European part of the USSR from 30% to 7.2% i.e. by about four times (Danilov, 1963). Considering this, and the utility of the polecat in destroying harmful rodents in fields and settlements, attention must be paid to husbandry of the stock of this carnivore. Damage caused by it to poultry is overestimated and can be easily brought to a
lated

by a decrease

in its

In the years after the

minimum

with proper construction of poultry yards.

1133

The

forest polecat

is

hunted chiefly

in late

autumn and

in the

beginning of winter by guns and dog, as well as with jaw traps

and various wooden snares or traps (cherkan, plashki and others). Hunting with dogs is the most efficient. In a season, the hunter rarely catch more than 10-15 polecats. The animal does not serve
as an essential element in
part incidentally captured.
It is

commercial hunting, and for the most

necessary to restrict the hunting of the forest polecat to

its

the period of complete "primeness" of

skin,

and not

to allow its

extirpation in the hunting season (P.Yu.).

STEPPE,

OR WHITE, POLECAT

Mustela (Putorius) eversmanni Lesson, 1827

1827. Mustela eversmanni Lesson. Man. de mammal.,
p.

144. Re-

gion of Khobda river depression in Ilek, south of Orenburg

(Stroganov, 1962).
1849. Putorius larvatus Hodgson. Journ. As. Soc. Bengal, 18, 447. Utsang in southern Tibet, north of Sikkim.
p.

1910. Putorius eversmanni Lesson var. michnoi. Kastschenko.

Kashchenko. Ezhegodn. Zool. muzeya Ak. Nauk, 15, p. 271. Kiran river 20 km from Troitskosavsk (Kyakhta), TransBaikaliya.
754

1912. Putorius eversmanni sibiricus Kastschenko. Kashchenko.

Ezhegodn. Zoolog. muzeya Ak. Nauk,

steppe,

17, p. 395.

Aginsk

Trans-Baikaliya.

putorius sibiricus

Nomen nudum. Nee Mustela Radde (= Mustela sibirica Pallas). Name

given with authorship of Radde (1862)^.

'"Kashchenko' s uses of the cited name is evidently a misunderstanding. p. 42), while describing the Transbaikal polecat which he called Mustela putorius and referring to its characteristics, emphasizes that its guard hairs are not black but as reddish as those of "Mustela putr. sibirica" as Radde writes. It is quite clear that Radde had in mind the kolonok, Mustela sibirica, but the addition of the incomplete word "putr." is a simple slip of the pen, a strange form appearing in this context. Therefore, to consider that Radde described a special form of polecat as is accepted by Kashchenko, has no foundation at all. Moreover, in the same text of Radde, there is nothing all that can be interpreted as a description of a new form. It is very strange that Kashchenko, after describing one form of polecat (michnoi) from Troitskosavsk (Kyakhta), noted another for the Aginsk steppe, i.e. from the nearest neighborhood. In the work of Kashchenko, there is nothing resembling a diagnosis. Apparently, he had a winter individual in one case, and in the other a summer, or an unmolted one.

Radde (1862,

1134
1913. Mustela lineiventer. Hollister. Proc. Biol. Soc. Washington,
26, p. 2. Chagan-Burgazy, Chuisk steppe, southern Altai.

1928. Putorius eversmanni satunini. Migulin. Zh. "Ukrainsk.

mislivets ta ribalka", No. 9, p. 30. Nogaisk steppe, north-

eastern Cis-Caucasus.

1927*. Mustela eversmanni robusta Ehik. "Nimrod", 10. Hungary. 1928. Putorius eversmanni talassicus. Ognev. Memuary Zool. otd.

Obshch,

lyubit.

estestv., antropologii

etnogr., 2, p. 26.

Talassk Alatau.
1928. Mustela eversmanni hungarica Ehik. Ann. Mus. Hist. Nat.

Hung., 25,

p. 37.

Mad'yarovar, Hungary.

1929. Putorius eversmanni occidentalis. Brauner. Zh. "Ukrainsk.

mislivets ta ribalka", No. 2-3, p. 9. Khersonsk gov.

1930. Putorius eversmanni amurensis. Ognev. Zh. "Okhotnik", No.

Blagoveshchensk on Amur region. 1936. Putorius putorius aureus. Pocock. Proc. Zool. Soc. London,
11, p. 25.
p.

703, Kazan.^'

1958. Putorius eversmanni nobilis. Stroganov. Izv. Sibirsk. otd.

AN

SSSR,

11, p. 150. Vic. of

Kokchetav, Kazakhstan.
p.

1958. Putorius eversmanni pallidus. Stroganov. Ibidem,

Kargat, between Novosibirsk and Lake Ghana.

150.

1958. Putorius eversmanni tuvinicus Stroganov. Ibidem,

p.

152.

Chaa-Khol'

village,

Tuvinsk depression.

1958. Putorius eversmanni dauricus. Stroganov. Ibidem, p. 154.

Smolenskoe

village, near Chita, Trans-Baikaliya.

1960. Putorius eversmanni heptapotamicus. Stroganov. Tr. Biolog. SSSR, 6.** Hi river, southern inst. Sibirsk. otd.

AN

Pribalkhash'e (V.H.).

Diagnosis
General color
light,

whitish-yellow, only weakly overlaid with

hairs).

blackish-brown (dark guard

Middle of belly

light,

some-

*Out of chronological order in Russian original Sci. Ed. "Belongs to the steppe polecat group. The place where the type was captured is, the individual was received by apparently, not accurately reported, or is incorrect

Museum from Latast, i.e. captured in those years when the steppe polecat was absent near Kazan. As can be determined by the description and drawing, it is possible that the type (the only specimen) is itself a hybrid of the black and steppe
the British

polecats (V.H.).

**Page number not given

in

Russian original

Sci. Ed.

1135
times with narrow, dark, longitudinal medial band, lower surface

of neck, chest, posterior part of

abdomen and
marked

inguinal region

brown

in color.

Base of

tail light,

only terminal part dark-brown.

Postorbital constriction of skull well

interorbital (V.H.).

narrower

than

755

Description

The white polecat

is

entirely similar to the black in general appearits

ance, proportions and habits, but

body appears somewhat more

hairs, especially those

elongated. Possibly, this

is

due to guard

on

the posterior part of the body, being not so long and luxuriant.

The

tail

is

quite short and constitutes about one-third of

body
is

length, covered by short close-lying hairs, and on the whole,

not very fluffy and appears quite thin.

Fig. 270. White, or steppe, polecat,

Mustela (Putorius) eversmanni Less. Sketch by A.N. Komarov.

1136

Winter fur

tall

and

soft, as a rule,

with dense short underfur

characteristic of polecats and sparse long guard hairs.

Length

in

both one and the other, however,

than forest polecat. Also less
is

is

usually less in steppe polecat

general thickness of pelage layer.

On

the whole, fur of steppe polecat

est polecat.

Guard

hairs

posterior portion of dorsal

is somewhat coarser than formore strongly developed and denser in region. At the same time, they are sparser

than in forest polecat and contrary to the case found in that

species, guard hairs never completely cover the underfur,

whose

color determines the animal's color on the entire body.

Anal glands well developed,

direct information on this).

their secretion has

same sharp

smell as in black polecat and, apparently, also can be sprayed (no

Teats, 3-5, usually 4 pairs.

General basic color tone of winter fur very

whitish-yellowish,

light,

yellowish or

background,

is

determined by underfur color. Over this a more or less significant dark frosting formed by

blackish-brown or brown ends of guard hairs (their bases are light). This frosting is stronger in middle and especially posterior parts of

where guard hairs are denser and longer, having a longer dark part, and it is more intensively colored. In anterior part of back, in the region of shoulder blade and along upper neck, this frosting, forming a sort of "two-layered" fur color, is weaker and may be almost undeveloped or even absent. Guard hairs here are extremely short. On the sides of the body dark frosting is also considerably more weakly developed than dorsally. Head piebald in color chin, lips and cheeks white; eye region, expanse between them and dorsal side of nose covered by brownish field (mask). Posterior to mask, crossing head from cheek to cheek runs a white band; in front of each ear, a small dark area of the same color of the mask is usually located. Occiput, region between ears and dark part of head usually has color of upper neck, on the whole forming dark area on head demarcating
the back,

posteriorly the white transverse band. Ears completely white, throat

yellowish-whitish or almost white. Character of head pattern very

756 variable both in color

and

in area of different color field.

is

Mask
light.

often narrow and pale, dark region

light

and whole head

Sometimes, head is entirely white or almost white and head pattern is absent.

(slightly ocherous)

1137
Lateral parts of neck of

same color

as anterior part of sides

lower surface, posterior to throat dark, blackishbrown or brown. Chest and forelegs black or blackish-brown;
or lighter;
its

Venter

light,

yellowish-straw;

its

posteriormost part, inguinal area

and posterior extremities dark, black-brown

Basal part of
its tail

as in fore limbs.

(half or two-thirds) has light color tone like

that of sides; distal part

brown or black-brown, almost

black.

Vibrissae black and

fitting

white; hairs covering soles of feet closedigital callosities

and

elastic,

black-brown and hiding

[in color].

and

foot pads.

Claws light-horn
is

Fur color

subject to very great variation, first of

all

being

individual. Its general tone

may be

either lighter or less light,

sometimes almost white, or with greater or lesser ocherous tinge. Differences in density and intensity of guard hair color exist, as do, consequently, intensiveness of dark frosting on basal light "deep layer" of color. Degree of development of guard hairs and darkening in the anterior part of body, it is generally said, are weak, sometimes disappear, and then, not only head but also the neck
and even anterior part of trunk are white or almost white. Intensity of dark fields on ventral body and, to a lesser extent, on tip of tail,
is

subjected to significant individual variability. Often, there

line

is

median

on the venter, a narrow dark

stripe extending

between
is

dark areas on chest and groin.

Summer
oped.

coat shorter and coarser than winter, and

not as

dense and close-fitting. Ocherous or reddish tone strongly devel-

On

head, especially around eyes,

brown tone more strongly

developed.

On

the whole, head darker than in winter; contrast

between dark and white sections sharper and it appears more piebald. As in winter fur, color contrast on head is sharper in younger
animals. In other respects, sumrrier color corresponds to winter.

Sexual differences

in color are absent.

first

Newborn

polecats are naked, but in

days of

life

they begin

to acquire, as in black polecat, a pelage of short

white hairs. At

approximately 10 days of age, limbs, forehead and back begin to

darken, and then facial

mask becomes

distinguishable and the animal

gradually receives a coat similar to adults, but duller, and formed by "underfur-like" hairs. Its full development of head pattern, i.e., dark mask, light (white) cheeks, white band behind mask and dark crown and forehead, is attained only in first winter pelage.

1138
In steppe polecat, the process of pattern
earlier
it is

development begins and proceeds more quickly than in the forest polecat, and already fully developed in the subadult individuals (Kratochvil,

1962). In polecats of intermediate age,

tern are well-manifested

they are

mask and

entire

head

pat-

relatively clearly outlined

and and

densely colored. With age, the whole head lightens, parietal and
occipital parts disappear or

become

paler, their area decreases

the

mask tone

lightens. In the very old polecats,

head

is

perhaps

entirely white, or

even silver

in tone,

and traces of mask are

unnoticeable or hardly developed.

As

noted, anterior part of trunk

ens,

and neck strongly lighten. With age, general tone of underfur lightwhich may lead to almost white [pelage] with light straw"Exotypic" variations which are so sharp
in

colored frosting.

black polecat are

not noted in the steppe, or they are rare. In particular mutations

like the "braginsk" polecat

have not been described.

observed
in

Geographic variation
black polecat,
757
is

in color, in contrast to that

better manifested,

intensity of general tone of underfur

and reveals itself in degree of (it may be almost white) and

guard hairs, degree of development of ocherous and reddish tones, density of dark ventral color, head and neck color, development of
facial pattern, length of black tip of tail, etc. Therefore,

well-

known
ity)

parallelisms of not only individual and age variation, but

is

also geographic variation

manifested. Density and length (qual-

of fur also changes geographically.

Skull coarse and heavy

heavier

and more massive than

in

forest polecat.

It is

relatively wide, with wider braincase

and more

widely separated zygomatic arches, and on the whole appears shorter and wider than black polecat, especially in braincase region. Mastoid width usually considerably greater than half condylobasal length

of skull. Protuberances, crests, etc. of skull more strongly and

sharply developed than in forest polecat, especially occipital crest,

and also

sagittal.

Also well developed

is its

anterior branch, ex-

tending to supraorbital processes, which, as in the mastoids sharply

expressed. Zygomatic arches strong and particularly widely separated in posterior portion. Zygomatic width in adult and old indi-

viduals usually exceeds mastoid width. Interorbital area forms clear

convexity on line of upper skull profile, from which profile line of

facial portion quite abruptly

descends downwards.

On

the whole.

1139

skull

somewhat

flattened, but in posterior region less so than in

black polecat.
Facial part of skull
is

not large, but

is

relatively larger

and

stronger than that in black polecat. Interzygomatic part of cranium

very narrow and extended. Lateral outlines of

constriction (interception; "waist") and behind

its

postorbital part

not parallel, but form two lines which meet at an angle in a sharp
it,

again diverge.

Width of
width

this part in its

narrowest place

is less

than interorbital
constricted part

in old individuals considerably less.

Most

in postorbital area,

with closed jaws, usually

lies in front

of line

uniting apices of coronary processes (see the note on page 1113).

External nares not compressed laterally; their width usually equal to their height. Auditory bullae in posterior half not swollen or

swollen to lesser degree than in forest polecat; carotid foramen lies in middle of longitudinal inner border of auditory bulla. Ends of
pterygoidal processes do not form strongly recurved external hook.

Nasal bones wide anteriorly, posteriorly narrowing to a quite pointed

wedge extending between

anterior portions of frontals, and, in

general, similar to black polecat. Dentition similar to black polecat

but stronger and teeth, most of

generally

all

canine and carnassial teeth,

more massive. In connection with this, lower jaw also somewhat more massive. Female skull differs from skull of male (not to mention measurements) by less sharply defined protuberances, crests, etc., and in having somewhat smoother outlines. It is lighter, with somewhat

weaker teeth. In young polecats, postorbital constriction not sharply defined; the younger the animal, the weaker. Moreover, it lies more anterior, closer to supraorbital processes. Braincase more swollen and
narrower posteriorly, without crests or only just noticeable. In connection with relatively wide interzygomatic area, whole braincase appears elongated and swollen. Skull of young steppe polecat
very similar to skull of black polecat of same age. Some features of this similarity are retained in older animals but there is another

age correlation
to that of a

the

skull of the subadult steppe polecat is similar

in

more mature black polecat

postorbital

some

respects (chiefly,

structure of interzygomatic area). Further, in connection with constriction

of

area

and

development

in

it

of

an "interception" species differences are projected all the more sharply. The course of age variation in the skull of steppe polecat.

1140

^g^.J??P^i
758
Fig. 271. Skull of steppe, or white, polecat,

Mustela (Putorius) eversmanni Less.

1141
759 therefore, in a purely

morphological sense,

is

considerably greater

than in the forest polecat. With time, apparently, such an essential

difference
is

absent, or

is

not proportional to the degree of mor-

phological differences. Apparently, the tempo of age changes in

the steppe polecat
is

trast in skull structure

more rapid. In one way or another, the conof young and extremely old steppe polecats
is

(with almost obliterated teeth)

much

greater than that in black

polecats of their respective ages.

As
skull

a very rare individual structural deviation, there

is

in the

sometimes a bony connection between the end of the pterygoidal process and the bony auditory bulla. The OS penis shows the same characteristics as in the black polecat, but differs somewhat in measurements (see below). The amplitude of variation of measurements of the steppe
polecat
is

very great and

is,

is,

in general, greater

than in the black

polecat. This

apparently, associated not only with the longer

growth period

in the steppe polecat, but also with

some geographic

which does not occur, or is less marked in, the black polecat. Moreover, the range of the white polecat within the boundaries of our country include regions where exploitation is practically undeveloped, and the animals live to the end of life, or in all events, grow to their maximum limit. The black
variation in this character,

polecat

is

hunted quite intensively everywhere.

black polecats, giant individuals are encountered

As
its

in

among

white polecats, the measurements of which extend beyond the limof "normal" fluctuating variation in the species. They are met
at least in
all

with significantly more often,

some
in

places, than the

black polecat. They are most of

found

western Siberia^^.

No

literature exists with information about giant polecats in other parts

of the range. Since unusually large kolonok (see page 1057) some-

times bear polecat features (dark-brown

I.M. Zalesskii, 1930),
it is

tail

end and brown

feet;

not excluded that the giant polecats (as

well as kolonok) are intraspecific crosses (in western Siberia, both

species live together and often in one biotope) and their size
is

an

expression of the phenomenon of heterosis. Giant polecats (as also

kolonok) are known almost only from their hides, and are nearly
unstudied.
along the upper Irtysh, they are even known to the local inhababnormally large polecats "mogil'shchik" (=grave digger) because of the belief that they live in graveyards and feed on the dead (Zverev, 1931).
^^In the steppes

itants

who

call the

1142
Sexual dimorphism in dimensions
length consitutes about
is

quite sharp

female body

85-90%
is

of male, and weight averages about

320-562 mm, of females, 290-520 80-183 mm, of females, 70-180 mm, length of hind foot of males is 40-80 mm, of females, 35-72 mm (from 1611 specimens: 1,237 specimens 671 males and 566 females from Zverev, 1931; 374 specimens from Stroganov, 1962); length of ear of males is 23-26 mm, of females, 20-23 mm
of males

45-50% of male Body length

weight.

mm;

tail

length of males

is

(Stroganov, 1962).

Condylobasal length of male skull is 61.7-82.2 mm, of mm; zygomatic width of males is 30.0-58.9 mm, of females, 30.0-48.7 mm; mastoid width of males is 35.747.9 mm, of females, 35.3-43.2 mm; interorbital width of males is 15.9-24.2 mm, of females, 14.0-19.5 mm., postorbital width of males is 12.0-17.2 mm, of females, 11.3-15.3 mm (Stroganov, 1962; values given by Ognev, 1931 and Novikov, 1956, do not cover the full amplitude of variation of body and skull dimensions). Os penis length of adult males (7) is 36. 7-M39. 1-42.5 mm, of young (30), 33.9-M36.6-40.6 mm; weight of bone in adults is 0.370-M0.436-0.530 gm, of young, 0.130-M0.200-0.280 mm (V. Popov, 1943; material from Povol'zhe). Even with some transgression in length, no transgression in weight of both age groups is
females, 52.4-76.7

observed).

Weight of Siberian males up

to

2,050 gm, of females, 1,350

tail

gm
760

(Stroganov, 1962).

One

giant polecat had a

body length without

of about

75-80 cm, although normal dimensions of animals from this region (in particular Semipalatinsk) is about 40 cm, and in rare cases 42-45 cm. It was taken in Semirech'e. Its color was completely typical for southern Siberian steppe polecats, but its fur was coarser and its skin "extraordinarily thick" (Zverev, 1931) (V.H.).
Systematic Position

The steppe polecat is undoubtedly very close to the forest polecat. The proposal to unite them into one species (Pocock, 1936) was supported by many, although not all, West European zoologists
having
rial

at their disposal,

genally speaking, only insignificant mate-

of the white polecat. The idea was not met sympathetically by

Actually,
if

us.

all

features and properties of both polecats are


1143
evaluated, and the question

is

approached not only from a purely

it

morphological point of view,

is

evidently correct to consider

them

different species.

Even

if

we

consider that the correlation of

these two forms (a group of forms) itself represents a "borderline

case" between species and subspecies in which either view might

seem

right, nevertheless in final consideration, the large

favor the treatment employed here

M.

(P.) putorius

argument and M. (P.)

eversmannP^.

Concerning skull structure, the white polecat

itself represents

a farther step of specialization in the direction of carnivory as

compared

to the black.

The white polecat has stronger

is

dentition,

protuberances, crests,

etc. are better

("predatory") musculature

developed and the masticatory much more powerful. Concerning this,

one can judge from the whole structure of the sharply compressed postorbital region indicating the degree of development of the corresponding muscles. Possessing no essential differences in body
structure, the white polecat as a whole,

and

in several of its indiIt

vidual races,

is

larger than the black polecat.

is

naturally as-

sumed
in

that all of this is related to larger prey.

The close connection

It is

of this polecat with ground squirrels and apparently even marmots

some places
is

is

better recognized

and understood.

associated

with murid rodents to a lesser extent than the black polecat and,
apparently,

completely independent of amphibians.

in

The course of age changes

between them
is

both species shows that their

skulls are very similar in early age

in the

no

essential differences exist

interzygomatic region. Later, in the black

polecat, the above-described structure of the interzygomatic region

formed, and

at this

age changes

in this region terminate. in the steppe polecat,

As

mentioned, these changes go farther

significant changes appearing,
all

with

strengthening with age, and lead-

ing to sharp differentiation from the structure characterizing the

black polecat. In very old individuals, the general appearance of

the skull has
little in

common

with the skull of the black polecat

less than with the skull of

mink. Therefore, the chain of age changes

in the white polecat is longer and leads to much greater differences between the skulls of old and young than in the black. Somewhat

schematically speaking, the skull of the adult black polecat corresponds with the subadult white polecat, and bears recogniz"For
details of the

whole question both morphological and geographical

sides,

see Heptner (1964).

1144
able infantile features. This relates even to old males, not to
tion females

menin

and males of younger ages

at the
is

same chronological
in

stage as white polecats.

The

correlation

approximately as

European and American mink, but the differences

of the final stages are greater.
for their treatment as independent species.

morphology

Geographic features of both species obviously give evidence However, if the entire
is

range of both species

761

taken into account, a definite geographical

vicariance

is

delineated, although the region of sympatry of the

great.

two species
belt
is

is

Together with

this, the

boundary "transitional"
if

not narrow, but a significant part

the range of the black

more of

about one-third polecat covered by the

is

not

range

of the white (the reverse correlation

exist in this region of mixing, in a

is

different).

Both species
species

known degree of independence,

inasmuch as there
itself to

is

the general possibility for

mammal

with this type of ecological isolation. The steppe polecat confines

open expanses, and the

forest

closed biotopes or human

USSR,
the black

settlements. In the southern European part of the

polecat penetrates into the steppe zone chiefly along the valleys of
rivers,
lives.

and

in the steppes,

with ground squirrels, the white polecat

[river], in its

Thus, along the Dnestr

lower course, the black

polecat lives in the valley, and nearby in the steppes, the white
polecat (Brauner, 1929); in the Carpathians, in the mountains

and to the south in the Carpathian plain the white (Konyukhovich, 1953), etc. Both species exist together in Czechoin steppeslovakia and Austria, but in different biotopes (the white type biotopes with ground squirrels) (Bauer, 1960; Kratochvil, 1962). They also colonized the Russian plain in the east and north
black,
(see later); the white polecat, as said, freely penetrated into the

the

region inhabited by the black. Therefore, there has been unlimited

contact between both species in the region of their sympatry of

inhabitation nonetheless. This

is

not contradictory in that in

floodland biotopes in some regions (Tatariya), their sympatry has

been recorded repeatedly (V.A. Popov). With all of this, black and white polecats give natural hybrids.
This
is

entirely natural

if

we

take into consideration the possibility

of hybrids of white polecat with kolonok and hybrids of the black

polecat with mink
polecat
is

forms the species independence

relative to the

undoubted. However, crosses of both polecats are of far lesser magnitude than one might expect from their sympatry and

1145
systematic proximity, and generally speaking, are quite rare. They have been noted in a series of places in the southern Ukraine, in Kursk and Voronezh districts, Trans-Carpathians and several other

places.

They

are

known

not only to furriers, but also zoologists^'*.

At the same time,

the

in their entire territory within the limits

of

USSR

(apparently, and outside the

polecats are encountered together, and

hybrid populations, and a

species
is

USSR) black and white nowhere do they produce zone of transgression between the two

absent.

Only

rare individual interspecific hybrids occur.

For Czechoslovakia and Austria they are not even recorded (Bauer, 1960; Kratochvil, 1962). In recent times, some western authors, on
the basis of their

own

observations in Central Europe, and consid-

ering the views of our taxonomists, recognize two species of

polecats (Bauer, 1960; Kratochvil, 1962)^5.

The North American

Bachon.)
is

ferret*

(M. (P.) nigripes Audub.

et

close to the steppe polecat of the Old World. Their

skulls are similar not only in

main

features, but also in all funda-

mental details (Pocock, 1936) including general dimensions and

The only difference lies in the color the ventral side (except legs) in M. (P.) nigripes is light. However, in several individuals a weak darkening is observed in
dimensions of the separate
parts.

the chest region, lower part of neck and inguinal region.

entirely likely, that
if

It

is

whole range of variation of all races of steppe polecats of Eurasia were evaluated, M. (P.) nigripes must be considered as only a subspecies of M. (P.) eversmanni. If we were to unite M. (P.) putorius and M. (P.) eversmanni in one species, then to give species independence to M. (P.) nigripes in no way permissible (V.H.).
the
762

Geographic Distribution

Found

in Central

and and

in part in

Middle Asia, Kazakhstan, southern

Siberia; southern

in part

middle parts of eastern Europe and

Central Europe.
possible
that

'It

is

one such hybrid, as far as can be judged by the

description and drawing of the skull, was used in the description of the form

Putorius putorius aureus from Kazan and was used as a reason to unite the species putorius and eversmanni in one species (Pocock, 1936).
see also the corresponding sections on black polecat,

"For the systematic interrelationships of polecats and related species of Eurasia, mink and kolonok. * The common name in English is black-footed ferret Sci. Ed.

1146

Geographic Range

in the

Soviet Union.

Range

is

great and constitutes a considerable part of the range of

namely its western and northern parts. The most westernly place of occurrence of the white polecat in the USSR lies in the Trans-Carpathian district, where it was recorded in the Uzhgorod and Mukachev regions (Kratochvil, 1962;
the species,

Konyukhovich, 1953). Apparently,

Carpathian plain, but
is

it

occupies the entire Trans-

absent in the mountains. This area of

is

from the remaining range of the species within the boundaries of our country and is connected with its central European part lying in Czechoslovakia,
occupation of the white polecat
isolated

Austria and Hungary. [This section] constitutes its northern edge, bordered on the north by the Carpathian [mountains]. Its union with the remaining range is made through Romania and the region

of the lower course of the Dnestr (see

later).

To the north of the Carpathians, the northern limit of the range of the steppe polecat begins in the Ravy-Russkaya region at the Polish border north and northeast of L'vov (Tatarinov, 1956).
Thence,
it

passes south around Lutsk and Rovno, going on to

Zhitomir and farther to Kiev (Sharleman', 1915) extending, evidently, somewhat to the north of it, and reaches Nezhin (Ognev,
1931; Sokur, 1960)
or,

more probably,

to

Chernigov.

It

is

not

excluded that in the expanse west of the Dnepr, the white polecat is locally distributed in areas lying to the north, although positive
data on this are absent, and in Byelorussia, this species
is

not

encountered (Serzhanin, 1961). Its spread to the north, going on in the more easterly parts of the range has not been noticed here, or
is

less intensive.

Farther to the east, in the expanse between the

Dnepr and

the

Ural mountains, the species being described had already spread

northwards quite long ago (see later), and especially intensively, apparently, in the last decade. Therefore, information on the northern limit of
its

range here

is

not always fully defined.

At the beginning of the 60's, it was accepted that from Chernigov the borderline steeply ascends to the northeast, directto ing itself towards Bryansk and even somewhat northwest of it

Zhukovka (Yurgenson, 1932). Thence, the border of the range

1147
extends almost directly eastward

to

Telech'e* to the north of

Orel (F.F. Rudnev) or even to Kaluga and from there to Tula,

the

Ryazan' and Gork'ii (Kuznetsov, 1952); i.e., approximately along Oka [river]. According older data, the occurrence of the steppe

polecat was recorded south of the line leading eastward from Orel,
at

north of

Chaplygin (Ranenburg), northwest of Tambov, at Morshansk Tambov (Vyazhlinskii, 1928) and at Kuznetsk (Ognev,

193 1)^^

Between Ryazan' and Gork'ii, the borderline takes a great bend to the north into Vladimir district. Here, in the beginning of the 60' s, in this region, the white polecat was met with everywhere, even the most northern regions of the district and was not recorded only in the very northwestern bordering Moscow, Yaroslav
and Ivanov
districts

western [regions]

Aleksandrov and one of the south Kurlovsk Vladimirsk Meshcher the west
in in
(in

to

of Shatura and northeast of Spas-Klepikov; according to the data of N.D. Sysoev). According to these data, the occurrence of the

white polecat in

Moscow

district is probable, especially since

later).

it is

not exclusively a rarity in Vladimirsk (see

It is

possible that the light [white] polecat penetrated into the

[in]

southern parts of Gork'ii

764 not only south of the Volga. In

Zavolzh'e and exists in this district any event it was recorded not only

for

Chuvash, but also

in the eastern part of

Mari

ASSR

(Yurgenson,
the

1932). Therefore, reaching the Volga

somewhere above

mouth

of Kama, the borderline directed

east,

itself quite steeply to the north-

covering not only the entire Tatarsk Republic, but also the
district.

southern part of Kirov

closer to

In an unclear

way

it

is

directed

Omutninsk (5840' N.
in the

lat. is

and a

little

east of 52 E. long.;

Popov and Lukin, 1949). This

by the species

the northernmost point inhabited

European part of the country.

*In Russian original, misspelled "Telyache"

Sci. Ed.
al.

'*Due to technical reason, the maps pubUshed by Heptner et

contained an error

(1950, 1956)

it

contained no data on the occurrence of the white polecat in

is

Moscow
In

district

and

absent in Smolensk. In agreement with the

new

data, the

border in the east extends in a different way.

some maps of

the range in the

European part of the

USSR

published before

the present time, their transgressions were very great (Bobrinskii, 1944; S.

Naumov

and Lavrov, 1948). This concerns also some other parts of the range, especially Priural'e, Priamur'e and Middle Asia, through which the data of the maps (Bobrinskii, 1944) in particular do not correspond with the actual situation.

l-i

cQ

1149

To
and

the east of the

European part of the

is is

USSR in

Priural'e

in the Urals, the

it

border of the range

apparently

very poorly clarified.

From Omutninsk,

directed towards the Urals (the

white polecat does not reach the northeastern parts of Perm district; Kuklin, 1951) and along its western foothills, and encompassing Tatarsk Republic from the east, which the white polecat
entirely occupies

(Popov and Lukin, 1949), steeply descends

to the

south. Southward, the border apparently extends approximately

along the Ufa and Belaya

[rivers]. At the latitude of Magnitogorsk, or slightly northward (53 30'-54*' 00'), the border turns sharply to

The assumption that the northern border of the range passes through the Urals below 58 N. lat.
the east and crosses the Urals.

(Yurgenson, 1932),

i.e.

that the steppe polecat lives all over the

middle and southern Urals,

In the Trans-Urals

is apparently wrong. and western Siberia, the border

at first

passes

northwards along the eastern slopes of the Urals through Miass (west of Chelyabinsk) to Sverdlovsk and, going farther to the north,
reaches approximately 60 N.
lat.

in

the upper Sos'va (Kuklin,

1938; Shvarts, Pavlinin and Danilov, 1951; Laptev, 1958; the north-

ernmost point inhabited in Asia). Along the Sos'va, the border goes to the southeast and reaches the middle and lower courses of the Tavda, and turning towards the northeast, it passes farther to
Tobol'sk or
to
to a point a little north of in
it.

But,

its

occurrence so far

the north

Trans-Urals sometimes gives rise to doubts

it

(Stroganov, 1962), and

has been assumed that the border of the

range from Sverdlovsk goes to

any event,
N.
lat.).

in

Irbit and thence even to Tobol'sk. In Trans-Urals the polecat does not reach Ivdel' (60 40'

From Tobol'sk,
or going a

the border line, generally following the Irtysh,

north, directs itself east approximately to the
it

little to its

mouth of the Shish river and there Vasyugan (I. Laptev, 1958), including,
of the

turns to the sources of

way, the entire basin (Yakushevich and Blagoveshchenskii, 1952). From the source of Vasyugan, it again extends to the east or, somewhat
in this
is

deviating to the north, and reaches the Ob' (the polecat

to the north of the

absent

Vasyugan basin) somewhere between the places

where Vasyugan and Ket' flow into it (I. Laptev, 1958). Farther, the border goes on to the Chichka-Yul river and crosses the upper Ket' and Kem' (V.N. Nageev) to reach the Yenisei between Krasnoyarsk and the mouth of the Angara, perhaps at Eniseisk
(Kuznetsov, 1952).

1150
Individual animals sometimes, apparently, go beyond the de-

scribed border, but long transgressions are very rarely seen. Thus,

was recorded at Narym (Ognev, 1931) and in 1947, one polecat was taken on the Ob' at Lokosov (Lokusov), somewhat above Surgut (I. Lapteev, 1958). If this is not a collected skin [from elsewhere], then it most probably is a traverse along the Ob' valley."
the polecat

To

the east, in the Yenisei basin (Irkutsk district), the northern

border of the range of the steppe polecat advances across the Shitka region to Biryus (a little north of 56 N. lat.) and Bratsk on the

Angara, and rising slightly to the northeast reaches Ust'-Kut on the source of the Lena. Thence, it somehow descends to [Lake]
Baikal without passing around
basin, there
765
is
it

from the north. For the Yenisei

at the

evidence of the capture of a polecat

conflu(at 64;

ence of the Viv' river with the Nizhni [Lower] Tungusk

Yakovlev, 1930; Ognev, 1931); this is a poorly understood mistake the range undoubtedly does not extend so far.

To

the east of Baikal, the range on the north includes the

left tributaries

region of the

of the Vitim, the Amalat, Tsipa and

Tsipikan rivers, and reaches the upper Angara (Bauntovsk and

Severo Baikalsk regions of Buryatsk ASSR; V.V. Timofeev,

Ananshkin). These are peripheral points where the polecat
rare and
is is

S.

very

very sporadically distributed. Thus, in Bauntovsk re-

gion,

it

occurs only in the upper course of the Tsipa in the region

of Lake

Okunevo

in

an area extending not more than 80-100

if

km

along the river. The polecat occupies a similar,

In the east,

not smaller, area

along the upper Angara near the mouth of the Kuta (V.V. Timofeev).

beyond the Vitim, the northern border of the range

sharply descends to the south and continues (Chitinsk district) across

the upper Chita river (Pavlov, 1948), then turns to the east and

across the middle course of the Nercha to reach the source of the

Amazar (somewhat south of Mogocha

mainline). Thence,
it

at the

[Siberian] railroad

steeply descends to the south to the Argun',

nearly in the region of the mouth of the Gazimur, and exits beyond

(80 E.

"Polecats were found at some points on the Ob' between the mouth of long.) and Surgut; i.e. north of 60 N. lat. were apparently, wrongly

map (1962) these were not clarified in the text and Old references (Kashchenko, 1900) to the distribution of polecats throughout the whole of "Tomsk Territory" are incorrect. North of Tomsk, the polecat does not pass beyond 57 N. lat. i.e. it occupies only the southern parts
placed on Stroganov's
it.

contradict

of

Tomsk

district.

1151
the frontier (Favorskii, 1936; P. Zimin). Therefore,
that the polecat

it

is

evident

does not reach the confluence of the Shilka and Argun', and contrary to the very widely distributed view, it does not occur in the upper reaches of the Amur.

A
is

separate small area inhabited by the polecat in our country

Amur. It occupies Blagoveshchensk and the lower Zeya and extends eastward from Blagoveshchensk to Chesnokov (80 km to the east) and Poyarkov, apparently reaching the Bureya and occupies its lower [course] and, possibly, the foothills of Bureinsk range (Radde, 1862; Ognev,
located on the middle

region

1931, 1935; L.G. Turova). This section of the polecat range itself represents the northern edge of the region inhabited by the species
in northeastern

China (former Manchuria) and

is

connected with
its

the Trans-Baikal region through this country. References to

occurrence along the

Uda (Radde,

1862; Ognev, 1931; Stroganov,

tially,

1962) are apparently, mistaken. This error may be, at least pardue to the fact that the data concerning the Uda a tributary

where the Siberian polecat is comof the Selenga (Baikal basin) mon, were discounted because of the Uda, flowing into the Sea of
Okhotsk.

From

the above-mentioned western point at Rava-Russkaya,

the southern border of the range of the steppe polecat directs itself
to the southeast, generally

going along the Carpathians through

L'vov and Stanislav, and farther envelops Chernovits from the north and east and somewhere in northern Moldavia, descends to the south and reaches the border of the USSR with Romania. Stepppe polecats are found throughout Moldavia (Kuznetsov, 1952) and in the region of the Danube mouth. In the Carpathians, this
species
is

absent; only the black polecat lives there.

Farther to the east, the range extends southwards to the shores

of the Black and

lives not only

Azov

seas,

and

in the

Crimea, the steppe polecat

on the plains but also in the montane part, but apparently avoids thick forests and is absent on the southern shore. In the Cis-Caucasus and Caucasus, the southern border of the range passes through the region a little south of Krasnodar along the foot of the northern slope through Nal'chik and Ordzhonikidze to Makhachkala on the Caspian Sea and thence descends to Derbent along the narrow coastal strip. In the west the border, approximately
at the

meridian of Gelendzhik, crosses the western spurs of

the [Caucasus] range and the [species] range extends southeast as

1152
a narrow strip along the shore itself nearly to Sukhumi.
cat does not ascend the

The

pole-

were wooded) and is m (Psebai) or even less (Dinnik, 1914; Vereshchagin, 1947, 1959; V.G. Heptner). Therefore, in the Main Caucasus and in the TransCaucasus, the polecat
the seacoast.
is

mountains (the foothills are wooded or known to a height of only about 600

absent. In the regions adjacent to the

northwest, north and northeast of the Caspian the range extends to

Precise borders of the steppe polecat range in Middle Asia
766

still

cannot be established. In any event,

it is

far

from being distributed

border of the range
exists bx)th in

everywhere here, and the

in this

line of the southern

country

is

quite complicated.

The polecat

Mangyshlak and on Buzach peninsula (Ognev, 1931; Gerbel'skii,

1939; Sludskii, 1953) and along the Caspian Sea (Kara-Bogaz-

M. Laptev, 1936) the range descends to Krasnovodsk as a narrow extension and, perhaps, may go along the Sea and a little
Gol;
farther to the south, although data about this are very indefinite. In

Bol'shoi Balkhan, the polecat

is

absent (V.G. Heptner, G.Sh.

Shukurov).

From Krasnovodsk, the border of the range, passing north around Bol'shoi Balkhan, extends towards the northeast and, apparently, along Ustyurt Chink in which the polecat exists, particuits eastern part (Bazhanov, 1951) reaching the lower Amu-Dar'ya. In the delta and adjacent parts, polecats were noted in Muinak, Kungrad, Takhtakupyr and KhodzheiF regions. To the

larly in

south, the polecat

is

distributed along the rivers, apparently, to

Khiva and Urgench (Gladkov and Nikol'skii, 1935; Kostin, 1959; 1961). The polecat is absent in all remaining areas of Middle Asia west of the Amu-Dar'ya, both in the Karakum and throughout the southern regions adjacent to Iran and Afghanistan despite several claims to the contrary (Bobrinskii, 1944 and
Ishunin,
others) (V.G. Heptner).

The data of the range of the polecat in the regions to the east of the Amu-Dar'ya are scarce and in part, not well defined. It exists in the northwest, directly adjacent to Amu-Darya delta, and in the northern part of the Kyzylkum desert here in particular, in the regions along the Kuvandar'ya and Dzhanydar'ya [rivers]

(Ishunin, 1961). In remaining part of the desert the polecat

ently, absent.

is,

appar-

No

clear positive data exist

and

that available, partial

and suggestive are not definitive (Kuznetsov, 1948; Sludskii, 1953).

1153
Farther, polecats exist, though sporadically distributed appar-

mountain system of the Tien Shan and in part, the Pamir- Alai. In the north, they are known in Dzhungarsk, Kirghizsk and Talassk ranges and in Karatau, and extend southward to the Alaisk valley. They are, however, absent in the Pamir (Rozanov, 1935; Flerov, 1935; Meklenburtsev, 1936). In the west, in Trans-Amu-Dar'ya part of Middle Asia, their occurrence was described between Chiili and Chimkent (Ognev, 1931), near Chinaz on the Syr-Dar'ya and in Dzhizaksk Golodnaya steppe (Sardoba;
ently, along the entire

Ognev, 1931; Obruchevo station in Za-aminsk region; T.A. Pavlenko), on the left bank of Syr-Dar'ya in Begouat region (east
of Dzhizak), in Khavast region in the foothills of the Turkestansk

range and

in

Samarkand and Bukhara oases (Shafrikan, 50

is

km

to

the northeast of Bukhara). Farther south, the polecat

the Karsha,

Guzar (southeast of Karsha) and Saryassii at Surkhandar'ya) regions (Salikhbaev, 1939; Dubinin, 1954; Meklenburtsov, 1958; Ishunin, 1961; T.A. Pavlenko; V.G. Heptner). Farther south and east, references to this species are absent
(Chernyshev, 1958).

known in (near Denau

Based on these places of occurrence, the border of the polecat in the Trans-Amu-Dar'ya part of Middle Asia can be, apparently, preliminarily be given thusly. From the lower Amu-Dar'ya, probably at the level of Khiva or somewhat below, the border line
range
includes parts of the desert directly adjacent to the delta (in particular, regions

of old cultivation) and then turns northeast, con-

tinuing along those parts directed towards the Aral Sea, and crossing

Dzhandar'ya and Kuvandar'ya districts, reaches the Syr-Dar'ya, somewhere in the Kzyl-orda district. Being directed towards Syr-Dar'ya or along this river, the border passes farther south, including the Golodnaya Steppe (at Dzhizak). Thence, it goes west along the Turkestansk range, probably south of Nuratau, where the polecat was not recorded (Meklenburtsev, 1937), and
the

probably,

includes the oases along the Zeravshan as far as Bukhara.

Apparently, from somewhere in the region between Samarkand and Bukhara, the border passes at first southward to Karsha, then turning southeast, it envelops the Baisun mountains (in Kugitangtau
polecat
is,

apparently, absent) and farther,

in a still

moving eastward, goes

it

on
767

to

Surkhandar'ya. Thence,

completely unclear form,

goes eastward across the Alaisk valley and exits to China. To the east of Surkhandar'ya, the polecat was not recorded (Chernyshev,

1154
1958). According to

some data

(Sludskii, 1953, map), the polecat

is,

occupies the whole Kzylkum, which

rect (T.Z. Zakhidov).

however, evidently incor-

Unique

in the

sense of

its

general outline, the range of the

its

species and, to a considerable degree,

sporadic distribution in

Middle Asia are determined by the fact that it avoids sandy deserts and in this country is a species to a great degree associated with the mountains and foothills, high montane valleys (Alaisk), and in
part with the elevated areas in the plains.

The sporadic

distribution

of the species
ditions, with

is

associated, probably, in addition to biotopic conthe polecat

some of our mountainous regions which may have penetrated from the east (Alaisk valley).
In the

whole expanse from the Alaisk valley

to the

lower Argun'

in Trans-Baikal,

the range of the steppe polecat extends south

beyond the border of the USSR. The high montane areas of the Altai and Sayan may constitute an exception, of which data, however, do not exist. The range of the white polecat has noticeably broadened westwards in the last century, and in part northward. However, the chronology of its movement was poorly traced. In Tatariya, even 100 years ago it was not present, evidently, and even in the beginning of the XIX century it was believed that it was absent along the western side of the Ural range (Eversmann, 1850). At the end of the 60's (1866-1869), M.N. Bogdahov did not record the white polecat in the "chernozem belt of the Povolzh'e" (former Simbirsk, Kazan, and Saratov governances, the eastern part of Penzensk and
it is not excluded that he did not always between the two species (he also did not refer to the white polecat for the lower Volga). At the beginning of the 90's of the previous century, the white polecat already inhabited all of former Saratov governance and in

western Ufimsk), though

differentiate

the beginning of our century penetrated into Tatariya but did not
still

occupy

it

entirely.

20' s, and this

was

the

species

known

at that

Near Kazan, it appeared at the end of the most northerly point of occurrence of the time (Ognev, 1931). In the Omutninsk reindications perhaps do not completely

gion of Kirov district the polecat apparently penetrated in the 40'

of our century.
If these

accurately describe the chronology of

movement of
is

this species to
it

the west and north in the Trans- Volga, in any event

can be

considered that the tempo of

its

colonization

high.


1155

In

middle Russia,

its

colonization northward also was local,

but the data concerning

this, particularly

on the penetration of the

polecat to the Oka, are insufficient and are not accurate (see above,

comparison of points of occurrence in expanse between Dnepr and Volga). In Vladimir district, where the described species do not represent a rarity (in 1960, 202 skins were prepared), it apparently appeared only at the end of the 30' s beginning of the 40' s, however it had reached the northernmost regions (N.D. Sysoev). Judging by the fact that it is absent in some of the westernmost

parts of the district (see above)

and

in

Moscow

district,

it

is

not
in

excluded that
part

its

settling proceeds not only

from the south but

from the
It

east.

apparently occupied Chernigovsk district mainly after the

20' s. In Ternopol' sk and L'vov districts, white polecats appeared

apparently, in the 40's of our century (first found in 1950).

The

stimulus for the colonization of the polecat here towards the west

was, apparently, the appearance of settlements of European ground

squirrels [Spermophilus suslicus] in this direction (Tatarinov, 1956).

The appearance of

the steppe polecat in Central

it

associated with recent times: in 1928

Europe is also was established in Hun-

gary (Ehik, 1928), after several years in Trans-Carpathia, in 1948 in Austria where it rapidly spread in Czechoslovakia and in 1952

and
768

its

number strongly increased (Bauer, 1952; Kratochvil, 1962;

difficult to establish the time

O. Wettstein)^^
It is

when

the polecat generally

began to move from Asia and settle in Europe. It went, apparently, mainly to the west along the south and as a secondary process from the south to the north. The latter movement, as was shown, is a phenomenon of our day and the last century. It is possible that its movement along the most southerly route, at least in some parts, began also in the recent past, some hundred
years ago. Concerning
ev. soergeli) are
all

of

this,

Pleistocene fossils of polecat (M.

known from

France, Baden, Wiirttemberg, Hun(Ekhik, 1928; Pidoplichko, 1951).

gary and Austria and the Pleistocene and Holocene from the southern European part of the

USSR

'4n

light

of the stated facts, information on the occurrence of steppe polecat near

is

Warsaw

(apparently in the last century; Ognev, 1931)

clearly wrong.

It is

possible

that the capture of a chromist black polecat, the so-called "P. p. stantschinskii"

was

the basis for this mistake.

penetrated into

Only a short time ago, the white polecat had Poland from Rava-Russkaya region.

slightly

1156

9^
V

"
30

80

90

120

10

V,

J^

^^^

180

~-'^ \
i_j2\
-.,-->
1

-li-^

( 1^

\^
4. *^
\

/r^i&SA V
St
iy<W--, Moscow^^^

M/ U^vI^V^^^'- *^^

lj^^^
^
'y'
)

^"^^^fe^^V^ \w
1

<^\

30

V ^ /\.p
<

^ \7
:'

^-

'"^"'^.

^^^
\
'

\
'

30

j U, (f^
1

'^

-*! )

/
~-^

p^^ A.
k7

^^[^^^^-^, -\^ N,>c^

-

'^

<^
JU

4-^ r7

J
1
/ /

\^< [y
30

.U-=wT
1

{7
20

i..J
150

60

90

768

Fig. 273.

Species range of the white polecat, Mustela (Putorius) eversmanni Less.

in

Southern and eastern borders

Asia to a considerable degree given provisionally.

V.G. Heptner.

Nevertheless, the present polecats of central Europe must not be

considered as autochthons. Obviously, the entire phenomenon, taken

on a large scale, must be considered as a secondary reoccupation

of the previous, for
If
it

some reason abandon

in the southeast
in

range.

is

now

difficult to say

anything about the time of the

appearance of the polecat

and

in part in central

Europe, then,

and the extreme south, any event the movement of

the northern, western and southwestern borders in recent times are

without doubt.

It

may be

partially defined as the settlement of the

1157
769 forest-Steppe. The fact that the range of the steppe polecat in Europe
is not yet completely formed is indicated by its very outlines, which do not completely correspond to the distribution of the natural

landscapes.

Some expansion
border there
is

of the range to the north

is

occurring locally

also in western Siberia; however, the progressive

movement of

the

very slow. In Siberia, colonization of polecat in

new

places

is,

apparently, directly associated with the cutting

taiga. In

down

and clearing of the

of the polecat in

Europe,

this

cause also has certain

places, colonization

significance (Trans-Kama); however, in

many
is

many

places

is,

apparently, determined by other,

apparently partially internal reasons. This

very obvious in the

forest-steppe and particularly in old cultivated regions in the

extreme west of the range, where conditions of existence have not changed for a long time.

The sporadic
Trans-Baikaliya

distribution of the polecat in Cis-Baikaliya and

among

separate isolated small steppe areas (Lena,

to think,

upper Angara and others) permits one

on the contrary,

was never large. some attempts were made to acclimatize the polecat somewhat more to the north of the border of its natural range (Narymsk district; Lavrov, 1946).
that the distribution of this species in Pribaikal

For the sake of accelerating colonization

in Siberia,

Geographic Range outside the Soviet Union

In Europe,
it

occupies Romania (apparently, except montane wooded

Hungary, western (Lower) Austria, Czechoslovakia, northern Bulgaria and northeastern Yugoslavia
parts of the Carpathians),
(Serbia).

Mongolian ReKashmir and Ladakh; in China Tibet and the eastern part of the country from former Manchuria (except eastern) in the north to Sichuan in the south, including parts of Inner Mongolia. The 770 polecat is absent in the Korean Peninsula. In this expanse in Asia,
In Asia, the range occupies Dzhungaria, the

public,

the range has, apparently, large lacunae for example, in the north-

ern taiga parts of the Mongolian Republic, they are mainly in the
vast deserts, and perhaps in high
erally, the

montane regions of

Tibet.
is

Genvery

range

in

Asia outside the borders of the

USSR

poorly known.

1158

769

Fig. 274.

Range of the American form of steppe polecat [black-footed ferret], Mustela (Putorius) nigripes Aud. et Bachm. (from Hall and Kelson, 1959, simplified).

closely related form, perhaps only a subspecies of our steppe

polecat (see above), M. (P.) nigripes Audub. et Bachin, inhabits North America, occupying a very small range located in an irregular strip

from southern Alberta and Saskatchewan in the north to Oklahoma, northern Texas and New Mexico in the south. It is very infrequent, apparently, endangered (extirpated), relict form (V.H.).

Geographic Variation
Despite
polecat

geographic variation of the white might have been expected, especially taking into consideration that the animal is met with in
the
vast

range,

is,

relatively, not as great as

1159
very different natural regions.
possible, that part of this

It

is

is

explained by the relatively recent colonization of the species in the western and most northern parts of its range (see above). Thus, it
is

noteworthy that variation

is

greater in the eastern and southeast-

ern half of the range.

In a morphological sense, geographic variation
is

of small

amplitude. The differences

lie in

changes

in general color (degree

of paleness of underfur, purity of dark color of the guard hairs and

degree of their development, and degree of development of ocherous

and red tones) and the color of separate parts of the body (head,
neck), different density and length of fur, color contrasts, develop-

ment (length) of the dark tail tip, and in part, in general dimensions (see above "Description" section and later).
Until a short time ago, geographic variation of our polecats,

and also polecats of China and the Mongolian Republic was very little studied (for the fauna of the USSR, only Ognev, 1931) and
in part, particularly, but unsatisfactorily, Central Asiatic polecats

(Pocock, 1936). Relatively few forms were, however, described,

and some of those were assigned to other species sometimes ("P. larvatus"); several from the very beginning were clearly unfounded. In 30' s 3 forms were recognized (Ognev, 1931) in our fauna (5 were described), and for the entire species 5 (Pocock, 1936, in-

cluding the form aureus). In recent years, a

new

revision of geo-

graphic variation of the polecats of Siberia and, in part, Kazakhstan

(Stroganov, 1958, 1960, 1962) led to the description of 5

new

subspecies, part of the time in obvious agreement with fur grades.

Therefore, a system of nine races was formed only for Siberia

(Stroganov, 1962) and of not less than 12 for the whole country.

Characters of this series of Siberian forms were based on

times extremely subtle differences, hardly perceptible in a series of

animals or even undetectable. ^^

The number of recognized forms

ation of the species.
is

is,

obviously, too great and

gives an erroneous and exaggerated impression of geographic vari-

Even

the

number of

the recognized fur grades

considered to be too large (Kuznetsov, 1952).

A
sary.

full

revision of geographic variation of the species

is

neces-

"The color plates in the work of Stroganov (1962) are technically so badly prepared that they cannot represent the individual color characteristics of the de-

may be completely distort them. Compare, for example, and descriptions of the forms lineiventer and tuvinicus.
scribed forms and

illustrations


1160

The enumeration below must be viewed as completely prelimigood and in part weak. Among them, three groups are more or less clearly recognized: 1) polecats
nary, of designated forms, in part

group of Europe, western Siberia and northern part of Kazakhstan Trans-Baikaliya and Priof hungarica-eversmanni; 2) polecats
Baikaliya, Mongolian Republic, Altai, mountains of Middle Asia

and Tibet
cats
771

group michnoi-larvata;

perhaps the less

known

pole-

of the plains of Middle Asia and southern Kazakhstan {''talassicd") may be related to this group, but more probably they

belong to the first group; 3) polecats of Priamur'e and the eastern amurensis. part of former Manchuria 1. European steppe polecat, M. (P.) ev. hungarica Ehik, 1928
(syn. occidentalis).

Dimensions small. Pelage sparse and coarse. Color relatively dark-brownish, underfur yellowish, strongly darkened by the darktawny tips of guard hairs. Neck grayish-yellow, tail half sandy
color, half dark-brown.

On
races.

the whole, color darker than in nominal

form and other

Information on dimensions absent. Westernmost part of range in European part of country, northwards apparently, to northern limit of distribution, eastwards, probably to Volga and northern Caucasus. Outside the USSR in Czechoslovakia, Austria, Hungary, northeastern Yugoslavia, northern Bulgaria, Romania and apparently in a small section of Poland at its boundary with L'vov dis-

trict.

There has been no direct comparison of materials from southern Russian polecats, described as occidentalis, with those of Central

Europe; however, their diagnoses are actually identical, and they

excluded that the range of

must, apparently, be considered identical. The possibility is not this form in our country is restricted to

only the Trans-Carpathian plains. Polecats of the eastern parts of the Russian plains and the Cis-Urals have not been analyzed from
the systematic side (see description of following form). Furriers

assign animals

from Astrakhan and Gur'ev

districts

to

the

"Russian grade", which evidently coincide with the described form.

2.

Petropavlov [steppe] polecat, M. (P.)

ev.

eversmanni

Lichtenstein, 1827 (syn. nobilis, pallida, heptapotamica, aurea).

1161

Dimensions moderate. Winter pelage dense and soft, of modLength of guard hairs 32-35 mm or 35-38 mm. General color tone pale, straw-whitish, sometimes with ocherous tinge; underfur white or slightly yellowish-straw, sometimes pure white. Apices of guard hairs pure black ("pitch black"), black-brown or brown. Guard hairs on posterior part of back almost fail to darken
erate length.

the basic light color tone or darken

it

little.

Neck

whitish. Tail

not heavily furred,

its

basal part mainly whitish, the distal third

black-brown.

Body
hind legs

length of males

is 320-460 mm, females, 280-410; tail 80-160 mm, females, 70-140 mm; length of of males is 40-70 mm, females, 30-54 mm (after Zverev,

length of males
is

1931 and Stroganov, 1962).

Average dimensions
(561) 390

(after

Zverev, 1931): body length of males

mm; tail length of males (519) mm, females (528) 110 mm; length of hind leg of males (566) 50 mm, females (664) 40 mm (material from southeastern part of western Siberia former Omsk and Slavgorod areas). Condylobasal length of male skull is 61.7-71.3 mm, females, 52.4-61.3 mm; zygomatic width of males is 30.0-47.0 mm, females, 30.0-40.0 mm; mastoid width of males is 35.7-40.0 mm, females, 30.0-35.2 mm; interorbital width of males is 16.8-20.2 mm, females, 14.0-17.2 mm; postorbital width of males is 12.016.6 mm, females, 11.3-15.2 mm (skull measurements after

mm;

females (671) 340

120

Stroganov, 1962; data of nominal and three other forms are

combined, excepting aured).

In Trans- Volzh'e, western Siberia east to Cis-Baikaliya (Irkutsk
district)

and south

to the Cis-Altai steppes, Pri-Balkhash, the plains

portion of Semirech'e (Alma-Atinsk district), and Kazakhstan,

except the southern part.

Outside the

USSR

[its

occurrence has] not been established.

It

may
772

possibly be encountered that part of China contiguous with

eastern Kazakhstan.
In different parts of
its

range, this

form exhibits

certain devia-

tions, for instance, in intensity of color of

guard hairs, and their

density; however, these populations hardly require separation into

separate races, although several have been separated. In Trans-Volzh'e, the described form evidently transgresses into the

European

Trans-Volzh'e polecats, for instance, the black tip occupies half the length of the tail, and the skin is quite darkened
in

1162

by the guard
defined.

hairs.

The

limit of the range in the east

is

also poorly

The form
Kazan,
is

"P. p. aureus" Pocock,

1936 described from near

not generally recognized, not only because of considera-

tions previously given, but also because this

form

is

apparently

based on a hybrid specimen (see above).

3.

Turkestan steppe polecat, M. (P.)

fur long

ev.

talassica

Ognev, 1928.

and dense, but somewhat without rusty tones; color very pale, whitish, coarse. General underfur slightly yellowish, guard hairs pitch-black, but length of

Dimensions small. Winter

black zone of guard hairs insignificant, so as to give a generally

pale color.

Head

whitish, and

mask hardly

noticeable, neck almost

white. Tail relatively heavily furred, basal part whitish, distal dark-

brown. Summer fur very pale, without reddish tones, dark ends of guard hairs weakly defined, and mask very pale.

Data on dimensions are absent. Southern edge of range of species, in area between Caspian Sea and [Lake] Balkhash the western borders of Tien Shan

(Talassk Alatau), southern Kazakhstan, Uzbekistan, Tadzhikistan

and Turkmenia. Absent outside the USSR.

very

little

known form,
It
is,

apparently occupying the plains of

belt of Kazakhstan, not penetrating

Middle Asia and the southern

high into the mountains.
Central Asian races (michnoi
is

larvata; Ognev,
is

possibly, related to the group of

1931); however,

it

evidently smaller and apparently

allied with those described

above.
In the fastness of the Pamiro-Alatisk

system (Alaisk valley),

later)

and possibly deep

in 4.

in the

Tien Shan another form exists (see

this

several respects, perhaps close to

form.

Baikal steppe polecat, M. (P.)

ev.

michnoi Kastschenko,

1910 (syn.

sibiricus, lineiventer, tuvinicus, dauricus).

large.

Dimensions very
hairs

Winter fur very long, somewhat coarse,

of back, individual ones even
its

with long shaggy guard hairs and sparse underfur. Length of guard

64-68 mm,

in posterior part

longer (up to 80 mm). Tail densely furred with coarse hairs,

terminal third black. General color pale, whitish, with a black guard hairs, sometimes with light reddish tinge. This is determined by

color of guard hairs, which are whitish at their bases and dark-

brown

or black at their tips and in the intermediate part, light

1163
reddish-tawny. Yellowish-brown

tint

is

more

distinct

on neck,

shoulders and back. Contrast between light underfur and long black
fur. Dark areas between the and hind limbs are aften united by a narrow dark band extending along the middle of the abdomen. Summer fur (Trans-

guard hairs creates sharply bicolored

fore-

Baikaliya) is very bright, rusty-reddish that depends upon the complete or almost complete absence of black color in guard hairs and brightness of underfur, which is comparatively long. Body length of males is 435-562 mm, females, 362-520 mm;
tail

length of males

is

of hind foot of males

of males
is

is

130-185 mm, females, 124-173 mm; length 54-80 mm, females, 48-72 mm; ear length

24.5-26.5

mm,

females, 23.5-26.0
is

mm. mm,
females,

Condylobasal length of male skull

63.0-76.7

64.6-82.2

zygomatic width of males is 37.3-58.9 mm, females, 36.9-48.7 mm; mastoid width of males is 35.7-47.9 mm, females, 35.3-43.2 mm; interorbital width of males is 16.2-24.2

mm;

mm;
773

females, 16.6-19.4

mm;

postorbital width of males

is

12.0-

(measurement from Stroganov, 1962; combined data of forms called by a number of synonyms). Cis-Baikaliya on the west to the range of the form eversmanni [in the east], Trans-Baikaliya, Tuva, montane parts of Altai espe17.0

mm,

females, 12.0-18.0

mm

cially the south

(Chuisk steppe).

Outside the

USSR western

part of northeastern

China (former

Manchuria), Mongolian Republic and possibly the northern parts

of Inner Mongolia and some (eastern) parts of Dzhungaria.
In

some
this

tions in color

about

form may show several deviaand measurements; however, really convincing data are lacking. It is possible that some populations have
places, the described
is

transitional characters with other forms. Thus, perhaps that

the
it

way

things stand in the northern parts of Trans-Baikaliya, where

was shown comparatively (Stroganov, 1962) that color of polecats living there represents a transition to the form amurensis. At the same time, to recognize, as was assumed by Stroganov (1962), the
presence of four independent races of polecat in that region, has

no foundation. The features recorded for them are too indefinite and subtle and the ranges are too small for a predator of that type.
5.

Tibetan steppe polecat, M. (P.)

large.

ev.

larvata Hodgson*, 1849.

Dimensions very
guard hairs black

Color light

underfur
Red

almost white,

in color, but quite sparse.

color tones absent.

*In original text, misspelled

"Hodgon"

Sci. Ed.

1164

of male skull

There are no data about body dimensions, Condylobasal length is 75 mm, and apparently even more, and female is and may be more (material of Z[oological] M[useum up to 70

mm

of]

M[oscow]

U[niversity]).

In Alai valley.

ently,

tics

USSR in Tibet, Himalayas, Kashmir and, apparmontane parts of Kashgaria. This form is very little known both as regards its characterisand also its distribution. It belongs to the group of large races
Outside the

of innermost Asia, in particular michnoi.

To

this latter

it

is

un-

doubtedly close.

It

is

possible that polecats deep in the eastern

Tien Shan also belong to the Tibet form. Here [in this book], it is only conditionally recognized until there is a more complete revision of Central Asian polecats.
It is

possible that

all

polecats from

Tibet through the Pamir-Alai, Tien Shan, Altai and Mongolian

Republic
6.

to Pri-Baikaliya

themselves {michnoi) represent one form.

Amur
is

steppe polecat,

M.

(P.) ev.

amurensis Ognev, 1930.

Dimensions moderate, apparently

Pelage
soft,

less than in preceding form.

short, of equal length

over the entire skin very dense and

with guard hairs appearing only slightly above underfur. Length

of guard hairs about 20

mm.

General color of dorsal side bright

reddish-ocherous, almost without dark tones in the posterior part,

or brownish-reddish both on back and sides. Tail weakly furred

reddish-yellow color through almost

very short

its whole length. Dark tip 32-36 mm against 60 mm and more in other forms, and has a brownish color. Body length of males (5) is 340-410 mm; tail length is 100140 mm; length of hind foot is 49-58 mm; length of ear is 22-28

about

mm.
Condylobasal length of male skull
zygomatic width
is

(4)

is

64.1-66.4

mm;

23.5-24.4
is

mm;

interorbital width is 19.3-20.1

mm. Weight

of males (5)

575-800

gm

(material

from north-

eastern China; Shou, 1958).

On

left

bank of middle Amur.

Outside the

USSR eastern

part of former Manchuria,

and

possibly farther south.

The Amur polecat itself comprises a well distinguished race. Against the general background of geographic variation of the species,
it

separates itself most sharply from

fur, also to

all

others. This applies

tail

both to general color tone of

difference in

color;

1165

and

to

very characteristic structure of winter

fur.

Although as

pointed out, some characteristics of this form are revealed in gen774 era! fur tone of

animals from Trans-Baikaliya, the

Amur

polecat
It

differentiates itself

from

all

other races or groups of races.

its

may
this,

be considered a representative of
it

own group

of races. In

is,

apparently the only form.

Steppe polecats living outside borders of our country have been very poorly studied, and their independence needs confirmation for both their morphology and distribution. Usually, the following forms are recognized: 1) M. (P.) ev. tiarata Hollister, 1913 Inner Mongolia, Gansu, Shaanxi, Sichuan; 2) M. (P.) ev. admirata Pocock, 1936-Chhili (Hebei), ? Shaanxi, Ordos. The form tiarata is close to michnoi but possibly still differs from it; the existence of admirata as an independent race is very doubtful. The North American M. (P.) nigripes Audubon et Bachman may possibly belong to this species (see above) (V.H.).

Biology
Population. The steppe polecat
predator; however,
is its is

common,

locally an abundant

populations are unevenly distributed, which

associated with the disposition of suitable habitats and their

qualities,

mainly with respect to abundance of principal foods.

is

The polecat
pikas.

particularly

rodents are abundant

numerous where relatively large steppe ground squirrels and hamsters, etc. and

For certain parts of the range, the density of inhabited burrows

fluctuates within great limits; thus, in the Sal'sk steppes, density

ranges from 8.3 to 1000.0 per 1000 hectares; in the Tersk steppes

from 5.0 to 40.0; in the Stavropol' steppes, more often 100.0 and up to 2000.0. In the Daurian steppes, the harvest of this animal in nine localities of 3 km^ each was an average 45.0 per 1000 hectares, ranging from 14.0 to 113.0. For small individual sections, the concentration of inhabited burrows may reach 26 per
hectare, taking into account the character of the colony (steppe

ravines) (Sviridenko, 1935; Yurgenson, 1948). In western Siberia,

characteristic density
is

10 burrows per 1000 hectares; for former


1166

Nizhne-Volzhsk territory about 3.6. The steppe polecat is most numerous in the Cis-Caucasus and the Kazakhstan steppes between latitudes 48-52 N. lat.
Figures for fur tanning are generally representative of the
populations of this species in the
years, the average annual harvest

USSR:

in the

pre-revolutionary

was about 900 thousand individuals. During 1924/25-1929/30, it fluctuated from 464 thousand to 1,296 thousand individuals. The yield per 1000 hectares for the whole range averaged 0.79, for the former North Caucasus territory 1.56, and for former Nizhne-Volzhsk 4.1, for Kazakhstan

territory

0.87.

Habitat.
level

The steppe polecat everywhere

prefers open habitats

and foothill stepppes, fallow fields, pastures, slopes of meadows and ravines, and semideserts. It rarely inhabits cultivated areas, alpine and water meadows, herbaceous forest openings, and sections of desert landscapes covered by saxaul forests. In winter, it readily visits the frozen banks of steppe rivers and lakes. "In
each case, the ecological setting
will
in

which the steppe polecat

it

lives

correspond to that inhabited by one or another of those roits

dents species constituting

ficult to describe

food objects, and therfore

is dif-

it

is

variable" (Sviridenko, 1935).

The steppe

polecat ascends mountains to an altitude of 1860

in the northern

Caucasus, and to heights of 2000-2200

Kazakhstan.

in the

mountains of

Food. The steppe polecat

is

a typical carnivorous animal.

it

The

polecat cannot assimilate plant food and

is

absent in the diet

(Zverev, 1931). Invertebrates are encountered rarely in

foods, and only in individual sections of the range.
775

summer

In contrast to the forest polecat, in the steppe polecat diet not

mouse-like rodents but the larger steppe rodents: ground squirrels,

hamsters, pikas and others, are of decisive significance. Structural
characteristics of
its

masticatory system (see above)

is

associated

with

this.

Polecats cannot deal with such large rodents as adult

the

marmots and they only feed on

on dead bodies.

young and injured animals and

The

best conditions for polecats occur in those places densely

it feeds on them almost throughout the year, in warm periods seeking them on the surface and in autumn excavating their burrows, which the polecat does easily and quickly. Young polecats and females often enter

inhabited by ground squirrels. In these cases

1167
the burrows of ground squirrels without widening the passages. In

steppes where ground squirrels are few or are completely absent,

they are replaced by hamsters or pikas in the steppe polecat diet,

or on the banks of water bodies, by water voles. In

some places

(separate regions of western Siberia and Trans-Baikaliya), con-

sumption of ground squirrels bears a strictly seasonal character (Velizhanin, 1931). The close connection of the steppe polecat with ground squirrels is reflected by the distribution of its numbers, density,

and

in the character of daily

etc.

and seasonal

activity,

Mouse-like rodents mainly the various species of gray voles [Microtus] and hamsters play the role of permanent substitute food in winter in some parts of the range and in others in times of depression in numbers of large rodents.

absence of stable settlement,

Fish, domestic fowl and carrion

become of some

significance

for the polecat along the shores of rivers

ficient

and lakes, or with insufregular foods. Birds are of secondary importance in the

food of the steppe polecat.

rarely grouse.

Among

harvested birds,

it

attacks gray

partridge [Perdix sp.] and willow grouse [Lagopus lagopus], but

Amphibians and

reptiles

do not occur often

in its

food, and not everywhere.

Geographical (see Table 69) and stand-type variation in foods of polecat are well marked. In Kazakhstan (Sludskii, 1953), in alpine 776 meadows and in the mountains, it feeds on montane voles [Alticola],
relict [S. relictus]

and long-tailed ground squirrels

[S.

marmots,
regions

in forests

on

on hamsters, small
Volga-Kama

undulatus] and
in

voles and

wood mice
etc.

sandy

great gerbils

[Rhombomys opimus],

Even primary foods (occurrence not less than 14-16%) cannot be considered significantly stable in its winter diet. Over eight
winter seasons in
territory

(Grigor'ev and Teplov,

1939), voles were annually found in food analysis, water voles

only in four seasons, and ground squirrels and hamsters, fish and

amphibians

in various

combinations were encountered

is

in

only two

seasons out of eight (each). This

associated with fluctuations in

in nature (Grigor'ev

abundance of these groups of food resources and Teplov, 1939).

In captivity, adult polecats ate

55-150

90.6 gm, and when fed only on beef they an average of 165.8 gm. Young polecat age seven months an average of 145.5 gm daily and the age of months 155 gm. When fed
average
ate
at

gm

of food daily, on
ate

at

2.5

ground

squirrels, the daily requirement declined to 131

gm. Young

1168
775 Table 69. Geographic and seasonal variation in foods of steppe polecat (% occurrence; Grigor'ev and Teplov, 1939; Kozlov, 1931; Zverev and Klimov, 1931; Sludskii, 1953; Fetisov, 1936; Brom et al., 1948)

Type of food

Volga-Kama
territory

NizhneVolga
territory

Northern Western Kazakhstan Siberia

TransBaikaliya

Winter

Summer Winter

Winter

Winter Winter

Summer
13.2

Marmot
Ground squirrel Hamster
Pika

3.86

_
47.0
47.1

5.69

Water vole
Great jerboa

10.66

Hamsters Mouse-like
rodents

0.92

_ _ _

_ _ _ _
12.5

_ 76.3 _
6.8

_
1.49

0.26

23.3

10.07
2.45
9.8

76.2

1.04

15.4
2.2 4.4 4.4

13.2

0.26 4.32
13.6

5.2 6.8

59.4

52.9

31.3

Mole-voles
[Ellobius]

0.36
7.53
3.27
2.0

_
103
4.9

7.4 0.8 0.8

2.6

1.7

3.0

45
2.2

Birds

Harvested birds

Common
partridge

Domestic fowl Small birds

Frogs
Fish

0.73

5.8

0.36
4.4
5.5

118

0.8

Carrion
Insects

of data

29.1

23.7

_
38

1.16

0.15

3.0

0.45

0.07

0.22

1348

0.97

766

12.2

Amount

550

growing polecats, therefore, require more food than the adult per unit of live weight. Given 36 g of food, death due to starvation occurred on the 37th day, and on increasing the norm to 72 g, the polecat died on the 48th day. Therefore, the polecat eats in one day a quantity of food equaling 1/3 of its live weight; a norm of 1/6 its body weight does not maintain life functions. Young at the age of 30 days begin to tear at killed ground squirrels, though still nursing their mother. At the age of 40 days, they make independent attempts to
kill

ground

squirrels, but unsuccessfully. After 10

more days, they can kill very young ground squirrels, but are not able to manage older ones (Sviridenko, 1935). According to other data (Zverev, 1931), steppe polecats in captivity eat from 70-250

gm

per day.

Steppe polecats make food reserves that are often significant. In western Siberia, the following were found in burrows of steppe

1169
polecats: 1) 12
5

young ground

squirrels; 2) 13

ground

squirrels; 3)

ground

squirrels; 4) 5 hamsters (Zverev, 1931). In

Volga-Kama

and Teplov, 1939), stores contained: 1) 4 one hamster and 4 water voles; 3) 3 water voles and 2 gray voles. Probably, these stores that occur are not always
territory (Grigor'ev

vipers; 2)

utilized.

Home

range. The steppe polecats do not belong

animals with a sharply-defined

hold to a relatively settled

home
of

range. In the

among those warm time of

the year, especially in places rich in

ground

squirrels, old animals

mode

life until

they destroy those

found within an area a considerable distance from their burrows. They very quickly destroy ground squirrels within a radius 120 to
150
are

from
all

their burrows.

Young

polecats do not systematically

attack

inhabited burrows of ground squirrels successively; they

active, most often overnighting in the burrow of ground which they have eaten. Most settled are adult females nursing their litters. But even they, just after the young animals grow up, begin to roam together with the young animals (Sviridenko, 1935). Therefore, permanent home ranges are not well defined in steppe polecats and their outlines often change. The polecat lives in one home range for a few days up to 3-4 months.

more

squirrel

With available food

daily activity in

in
is

abundance and

shelters, the

range of

summer

small. In winter, steppe polecat

moves
in

12-18
it

km

per day, often returning to the same burrow from which

set out to hunt.

case as 3

The area of daily km^ (Zverev, 1931).

activity

was determined

one

In winter, with

worse conditions for capturing food, greater

depth of snow cover, the animals

to them.

move

to areas

In such case, significant aggregations are

more favorable sometimes

formed, especially along the slopes of the steppe ravines or near

settlements and

Kazakh winter camps (Kazakhstan; V.G. Heptner). Burrows and shelters. With rare exceptions, the steppe polecat

does not independently dig its burrow, but uses those of steppe rodents marmots, ground squirrels, hamsters, mole-voles, jerboas

777

and others,

after slightly

widening them. Due

to this, internal con-

struction of burrows occupied

sions, depth,

by steppe polecats,

their

dimen-

number of entrances, etc. vary greatly (Sviridenko, 1935; Zverev, 1931). Moving frequently, they are little concerned about a well-constructed burrow. They adapt to the burrows of their prey, whose mode of life is similar to these predators. In

1170

many

cases, the

burrow

is

only used once, sometimes for several

days; and after that not visited for weeks.

Nesting burrows characteristically possess

3 to 15-20); moreover, alongside the nest

many

outlets (from

chamber

for storing fresh food. Lining of the nest

is

chamber is a second chamber does

not occur. If one

present,

it

is

the remains of the nest of the

previous host. Characteristic signs of an inhabited burrow are pres-

ence of tracks, excrement, food remains and freshly thrown-out soil "shovelled" in 2-3 different directions. In burrows of males,
is 9-12 cm, in female bur5-6 cm, rarely 7 cm). In soft rows cm soil, the diameter of passages is larger. Most often, polecats use descending passages of ground squirrel burrows, which are usually wider than vertical ones. Independently dug burrows are usually shallow and simple they are used most often as temporary shelters in places where existing shelters are absent. Daily activity and behavior. The steppe polecat has no sharplydefined rhythm of daily activity. It is mainly a crepuscular-nocturnal animal, active most of all at twilight and at dawn. Often, particularly in the warm time of year, it is met with during the day.

the diameter of passages in hard soil

7-8

(in

ground

squirrels,

Apparently, the short

entire broods

summer

night

is

not sufficient for capturing

food, especially for females nourishing large broods. Hunting by

was observed during the day more than once. The influence of weather on the activity of the steppe polecat
is

has been professed, but

at

not strong. In winter,

in

it

is

decreased

30C

and lower. Daytime activity

summer

is

mainly ob-

served in cloudy, not very hot weather and in winter, mainly during snowstorms or in foggy, cloudy weather. According to data from laboratory investigations (Kalabukhov, 1943), the rhythm of

nocturnal activity in the steppe polecat

the forest polecats.

is

less well-defined than in

They

are considerably
all

more

active in daylight

hours.

From 22
to

to

55%

of

activity proceeds during the period

in

from 0600

1800 hours (36.5% on the average against 17.9%

the black polecat). Females are

more

active at night than males.

The steppe polecat is an active, clever, brave and curious aniIt swims excellently and climbs well, but rarely use these abilities. They dig out the burrows of ground squirrels very rapidly. It catches its food in burrows and on the surface by waiting and hiding. While hunting ground squirrels, the polecat does not
mal.

behave

in

a specific manner: after taking a certain direction,

it

1171

makes a great
attracts
it,

rush, often passing over inhabited burrows.

While

hunting, the polecat searches clefts, burrows and every object that

even

if it is

not experiencing hunger. In winter,

it

hunting mouse-like rodents,

the

snow (up
its

to

18-20

in

when makes numerous "plunges" under one hunt). Deep snow makes such
productivity, and the polecat
is

means of catching food of little

to

obliged

snowy one (Sviridenko, 1935). Usually, the polecat moves in large leaps, and in places with an abundance of prey, it makes frequent loops. In contrast to other small predators which plunge under the snow, it returns to the surface through the same "plunge hole" and not in another place. The polecat can overcome tetraonid birds in their snow roosts. It is only able to catch ducks injured by hunters.
change
habitat to a less

In connection with feeding to a great degree on large steppe burrowing rodents, the steppe polecat, as told, highly elaborated mobility and frequent changes in its home range. This did not help to develop a well-defined reflex for protecting its home range. The

steppe polecat almost alone

778 species forming, with [prey]

among mustelines

is

nearly the only

abundance and a high concentration of food, colonies of high density. This is assisted in that, under such conditions, broods of young polecats are often large, do not disperse for a long time, and hunt together in a quite small area. For this reason, young animals are observed with an absence of
aggression towards animals similar to them.

its

Seasonal migrations and transgressions. Quickly destroying food resources, and not experiencing lack of shelters, steppe

polecats, as a

measure of diminution of food

in a

given place,

migrate to another region of rodent abundance, sometimes moving

to

completely different surroundings (Sviridenko, 1935). They do

It

not live in one place for a long. time, especially the young.

was

also noted that steppe polecats colonized along the track of colo-

nizing ground squirrels. Thus, in the former Sal'sk province on the

left

bank of the Sai

river

where the ground

squirrels recently ap-

peared, steppe polecats were not previously

known

(the

same was

recorded in the steppes of North America where the same relationship exists between them* and prairie dogs, Seton-Thompson, 1921).
In 1948, in the Daurian steppes, an unusual concentration of adult

steppe polecats was observed in separate places as a result of local

migrations; in one case, 10 adult polecats were caught in a two*Black-footed ferret (Mustela nigripes)

Sci.

Ed.

1172

week period in May in an area of 3 km^; in another case in the same area, 33 polecats were caught in one month, 14 of which
were caught on three successive days. Winter weather leads
(deep snow, see above). In autumn, polecats
to

unfavorable conditions for obtaining food, causing such movements

move more

than

20

km

per day (Sludskii, 1953).

Reproduction.

Among

representatives of the mustelid family,

polecats are characterized by the most primitive type of seasonal

The period of complete sexual dormancy from September to January. In captivity, mating of steppe polecats was observed in the first third of March (Zverev, 1931) and at the end of the month (Sviridenko, 1935). In the Moscow zoo, mating (7 cases) was observed from 9 April to 9 June. External signs of estrus appeared on 12-13 March and developed for 2sexual cycle (Kler, 1941).
lasts

3 weaks. Then, after mating, they gradually disappeared within

3-4 days (Krumina, 1934).

In western Siberia, the period of rut occurs in

March. In the

northern Caucasus, mass mating takes place from the beginning of

February to the middle of March. Sexual activity of male polecats in Trans-Baikaliya was observed up to the end of May, and by the

end of June their testes had decreased in size to about 0.5 cm. The act of mating lasts from 20 minutes to 3 hours. In the absence of
productive mating, estrus
sive pregnancies. Usually,
is

repeated as
it

is

also the case after death

of a brood. In the latter case,

it

may be

repeated for three succes-

occurs only once, and the female does

not nourish more than one brood in the course of a year.

Pregnancy lasts 36-42 days (L. Vakhrameeva), or 40-43 days (Krumina, 1934). In other experiments (Sviridenko, 1935) in three cases pregnancy lasted 36 days after the first mating. Placentation occurs after 14 days, of which the latent phase lasts 7-8 days
(blastocyst stage; Kler, 1941).

The number of young in a litter is from 3-6 to 18. Average number from 33 cases recorded in the literature is 9.5. Growth, development and molt. Steppe polecats are born blind, naked and pale rose. Ears are covered by a membrane. Body length
is

6.5-7.0 cm, weight

is

4.5

gm

(Sviridenko, 1935). According to

data of

Moscow

body weight is number of young

Zoo, the body length of newborns is 5-6 cm and 10 gm. These figures fluctuate depending on the
in the litter

(Krumina, 1934).

1173
After three days, a thin white underfur appears on the body
surface.

On

the 9th day, the

increases six-fold (33 gm). At the

body length doubles and body weight same time, milk teeth begin to

erupt and the feet are observed to darken.

On

the 20th day, the

color darkens and typical difference in color of the muzzle appears.

At

that time, the

young polecats weigh 70-72 gm. Teeth

and

continue to erupt and color to darken. Eyes open on the 28th-34th

779 day.

At

that time, the polecats already crawl freely, lick blood,

even

try to tear apart prey,

although continuing to feed themselves

on the mother's milk. Body length of a one-month polecat is 190 and its weight is 138 gm. At the age of 45 days, they can already themselves deal with young ground squirrels and at the age with adults. They live in the family burrow for 2-2.5 of 60 days months. Dispersal begins from July, or later depending on time of delivery*. Young polecats living independently are already ob-

mm

served in September. Only the female takes care of the brood, but
in a series

of cases, participation of a male in feeding the brood

was observed.
Polecats attain sexual maturity at 10 months, but complete

development of the organism

is

reached only

at the

age of 2 years,

when skull formation is completed. The steppe polecat molts twice

tumn.

per year: in spring and in au-

Enemies, diseases, parasites, mortality, competitors, and population dynamics.

fox, corsac fox, imperial eagle
in the

The steppe polecat has many enemies: wolf, red and snowy owl. However, their role

population dynamics of the species is hardly significant. Competitors of polecat include: solongoi, kolonok, weasel, ermine, forest polecat, manul, steppe [red] fox, and corsac fox, as
well as
all

species of diurnal and nocturnal birds of prey living in

open

areas. In

summer, the polecat coexists

the ecological borders of
its

in contact

with the

majority of species of small carnivores, except for solongoi and

weasel, only
range.
at

distribution within the

Among
to rodent

infectious diseases, the polecat

is

weakly susceptible
disease

[sylvatic] plague, tularemia

and

viral

canine

distemper. Only weak individuals are susceptible

to pasteurellosis.

Helminthic infections are widespread among polecats. Ticks strongly

attack the steppe polecat.
It is

parasitized, often in great numbers,

Literally, "appearance in the light"

Sci. Ed.

1174

779

Fig. 275.

Paired footprints of steppe polecat while quickly jumping on snow, and

the track.

scheme of

Voronezh

district, 8 January 1952. Sketch by A.N. Formozov, about 2/3 's natural size.

by more than 11 flea species, partly transmitted rodents on which they feed. There are no data on mortality.

to polecats

from

Population fluctuations in steppe polecats are clearly evident.

They

780 both adult

by food deficiency, epizootics and the death of and young polecats as a result of steppe fires in dry years, unusually large flooding of steppes by snow-melt water in spring and also massive treatment of steppe lands with poisons for
are induced
the struggle against harmful rodents (Sludskii, 1953; Zalesskii, 1931;

1175
Sviridenko, 1935). In 1938, a catastrophic

fall in

number

all

over

Kazakhstan was observed as a result of the severe ice storms in early spring which hindered hunting by polecats, and following

which many rodents died (Sludskii, 1953).

In the steppes of southeastern Trans-Baikaliya, population fluc-

tuation of the steppe polecat

tions
is

is

well noted, but the range of fluctua-

less than that in other carnivorous animals in this region. In to the very

1939-1948, these fluctuations were 11 times higher due

high rise in their number in 1939, which was associated with a rare
gregarious vole [Microtus socialis]*.

abundance of the Daurian pika [Ochotona daurica] and groups of From 1940 to 1948, changes in

numbers were only four-fold. During these nine years, the changes in numbers of the steppe polecat were parallel to those of the longlegged buzzard [Buteo hemilasius] and almost coincided with changes in the numbers of Daurian pikas and narrow-skulled voles.
Field characteristics.
polecat in open habitats.
It is

not difficult to distinguish the steppe

Its

tracks on snow, sand or dirt are well

distinguished from tracks of solongoi, ermine and weasel by their

larger size.

Aid

in

determination of species attribution of the tracks

may always be

the habitat in which they occur, since in those where steppe polecat coexists with forest polecat or kolonok, they almost always occupy different habitats. The prints of claws and
callosities of the soles of the steppe polecat are

always very sharp;

In

the length of
it

its

leap

is

40-65 cm (Formozov, 1930).

moving,

often "trots" (P.Yu.).

Practical Significance

The

significance of the steppe polecat to the national

economy

of the

USSR

is

great.

It

is

particularly valued as a destroyer of

rodents harmful to agriculture, mainly ground squirrels. According

to very minimal calculation, the steppe polecat can destroy no fewer than 200-300 ground squirrels per year, not counting mice and voles.^ In regions where the steppe polecat feeds mainly on mouselike rodents, it can destroy about 1500 specimens in the winter alone (Sviridenko, 1935). In regions with weakly developed agriculture, the steppe polecat is very useful, destroying natural foci
is "stadnykh" Sci. Ed. an active season, one ground squirrel eats about 6 kg food, which consists of grains.

*The Russian word

*In

25%

of

1176
of disease transmitters and reservoirs where infections begin

ground squirrels and other steppe rodents.

The steppe polecat

is

very important for the fur trade of the

USSR;

in

individual years, the price of

seventh place; in

its skins has occupied Kazakhstan and several other places, the steppe

belt polecat occupies

one of the

first places. Its

durable beautiful

was always in demand in the national and international marHowever, in past years, from 1926-1929 to 1956-1959, the catch of steppe polecat noticeably decreased. Along the Cis-Cau6 times, in Midcasus, the catch decreased 4 times; in Bashkiria 14 times, 10 times, in the republics of Middle Asia dle Povolzh'e
skin
kets.

and in Nizhnyi Povolzh'e 17 times (Danilov, 1936). Apparently, besides profound changes in the economy of the fur trade, a considerable role is also played by such factors as changes in steppe landscape and decrease in food resources (ground squirhamsters and others) in connection with the application of chemical methods for control of rodents, with ploughing of virgin soil, and with changes in agrotechnical methods, etc.
rels,

Harvesting
781

this

animal

is

simple and profitable;

it

is

mainly

taken with jaw traps placed near the inhabited burrows. In the northern Caucasus, up to 100-150 were caught in one season us-

jaw traps (Sviridenko, 1935), in Nizhnyi Volga up to 60 polecats by one hunter. The best method for maintaining the number of steppe polecats at the desired level, which is usually determined by the oping 10-16

posing interests of fur trade and agriculture,

of
its

is

periodic prohibition
less

harvest for 2-3 years.

The prohibition must include not

it

than a whole district or territory, otherwise,

tive. In districts

will not
it

be effec-

of field-protecting forest belts,

is

necessary to

prohibit completely the harvest of steppe polecat during the whole

period of planting for the protection of the fields and the young
seedlings from ground squirrels (P.Yu.).

Genus of Marbled Polecats

Genus Vormela
Blasius, 1884

1884. Vormela. Blasius. Berichte d. Naturforsch. Ges. Bamberg,

13,
p. 9.

Mustela sarmatica Pallas = Mustela peregusna

edt.

Gueldensta


1177

General dimensions small.

Skull small, relatively short and wide, in general appearance
similar to skull of steppe polecat, Mustela (Putorius) eversmanni,

but smaller than

it.

oped

weak, but quite pronounced, anteriorly bifurcating and giving off branches to postoccipital well-defined, sagittal relatively
orbital processes. Brain case relatively small, not elongated, quite

Protuberances, crests etc. of skull well devel-

wide; zygomatic arches strong, quite sharply separated, only slightly wider than braincase in its widest place. Postorbital processes
quite large and massive.
In postorbital region a sharp constriction is present
lateral

line of
at

borders of this part of skull begin to

come

together

an

angle and then diverge again. Least width in region of postorbital

constriction less than interorbital width.
is

On

the whole, this region

relatively short. Skull noticeably flattened. Line of upper profile

It is

of skull relatively straight.

highest of

all in

occipital region

and gradually declines anteriorly; elevation in interorbital region is weakly defined; line of upper profile in facial part not reduced sharply. Diameter of infraorbital foramen considerably smaller than
canine alveolus.
lar triangular) in outline,

Auditory bullae moderately swollen, somewhat angular (irreguonly slightly more widely separated in

posterior part than in anterior.

On

the whole, bullae

seem

to

be

displaced forward and their anterior edge nearly reaches to the

foramen ovale, posterior ends of hook-like processes of pterygoid bones touch and are fused with anterior part of bullae. Mastoid

(mammary) processes
processes

well

relatively weak, paroccipital

(exoccipital)

developed.
3

Bony

palate wide.
1

Dental formula

I-C-P M = 34.

13

Teeth are strong

rela-

tively a little stronger than in

upper, are longer.

Main cusp of

genus Mustela. Canines, especially last lower premolar (third tooth

behind canine

in front of carnassial) has

no additional cusplets
is

on inner
width of

side.

Lower

carnassial tooth bears small additional cusplet

on inner side of main cusp. Upper carnassial tooth

its

strong
length.

cutting part approximately equal to half of

(last in the series)
is

its

Upper molar

has inner and outer sides of equal

782 length, or inner side

tion

constricted appearing as
is

slightly more massive last tooth is not two blades (inner and outer), or constric-

ill-defined.


1178

Trunk
length of

thin

and elongated, limbs

short, tail relatively

long (about half

and broad. Anal scent glands present. Fur relatively short and coarse, more or less even in length throughout the whole body; tail fluffy. Color entirely unique bright and variegated, consisting of patches and areas of black, yellow and white colors. There is no distinctive light throat patch. On facial surface of the head, well-marked black-and-white mask.
small, blunt; ears large

body plus head). Head

782

Fig. 276.

Reconstructed range of genus Vormela Bias., and reconstructed range of

marbled polecat, Vormela peregusna Gueld. V.G. Heptner.

1179

terrestrial animal, associated

with open, arid, level areas and

with burrows of mass settlement of rodent species (ground squirrels, gerbils).

specialized predator, feeding mainly on rodents.

Range of

the genus relatively small

and occupies southeastern

Europe, parts of Asia Minor, Middle Asia, Kazakhstan, and northern parts of Central Asia (for details see following description of
species range).
783

strictly characterized, and of independence no one raises any doubts. It long ago evidently even in the Pliocene constituted a group, with its origin probably
its

Genus is monotypic. Genus Vormela is very well and

in

Asia Minor

(fossils are

known from
its

the Pleistocene in Syria,

Palestine and the Caucasus). In

systematic position, the genus

apparently stands closest to the genus Mustela, namely, to that

branch which includes the true polecats (subgenus Putorius).

If

one of
side,
it

its
is

main features most similar to

its

entirely peculiar color

is set to

one

the steppe polecat.

Speaking essentially

of skull structure, actual fundamental differences are only in structure of auditory bullae

and

their location relative to the pterygoid

bones and the foramen ovale, and particularities of the dentition. In all remaining aspects, craniological differences between these two species are not greater, and in some respects, less, than those

between several species of the genus Mustela. Even color, with

of
its

all

peculiarity,

is

similar to the color of polecats in

some

char-

acteristics (dark venter, facial mask).

Genus Vormela has

several features of similarity particularly in

color, to different African forms of the family (Ictonyx, Poecilictis,

Poecilogale). Attempts to closely affiliate the genera Vormela and

Ictonyx (Pohle, 1933) did not meet with acceptance (Pocock, 1936),

although between them there are some

common

craniological fea-

tympanic bullae). Equally with genus Mustela, but particularly subgenus Putorius, genus Vormela, as was shown earlier, reveals characters with proximity to the monotypic North African genus Poecilicitis (libyca; Zorilla
tures (fusion of pterygoid processes with

libyca auct.) and is usually placed between them in the system. Sometimes, Vormela is considered more closely related to Poecilictis

The whole question of generic groups closely related to Mustela requires special study.
than to Putorius (Mensel, 1881).
there is one species: marbled polecat, Vormela peregusna Gueldenstaedt, 1770.
In composition of fauna of the

USSR

1180

Found

in

steppes and deserts of southern European part of

Asia, and Kazakhstan.

USSR, Causasus, Middle

fur-bearing species, also of significance as a regulator of

in

numbers of several rodents,

infections (V.H.).

particular reservoirs of harmful

MARBLED POLECAT, PEREVYAZKA'

Vormela peregusna Giildenstaedt, 1770
1770. Mustela peregusna. Giildenstaedt. Novi

Comm.

Acad. Sc.
d.

Imp. Petrop., 14,

Reiches.
river.

1,

p.

441. Steppes along lower Don.

1771. Mustela sarmatica Pallas. Reise durch versch. Prov.

1,

Russ.

p.

175, 453.

Between Syzran' and Medveditsa

1910. Vormela koshevnikovi. Satunin. Zool. Anz., 36, p. 59.

Ashkhabad.
1910. Vormela
784

tedshcenica.

Satunin.

Zool. Anz.,

36,

p.

60.

Tedzhensk oasis, Turkmeniya. 1910. Vormela sarmatica alpherakyi. Birula. Izhegodn. Zoolog. muzeya. Ak. Nauk, 15, p. 333. Near Ashkhabad. 1910. Vormela negans. Miller. Proc. U.S. Nat. Mus., 38, p. 385, plate 17. Ordos desert, about 100 miles north of Yulinfu
(Yuilin').

1935. Vormela peregusna peregusna natio intermedia. Ognev. Zveri

SSSR

pril. stran, 3, p.

70. Starogladovsk Station on Terek

[river].

1976. Vormela peregusna euxina. Pocock. Proc. Zool. Soc. London,

p.

718. Malsosi, Dobrudzha, Romania.

syriaca.

1936. Vormela peregusna

Tiberias* Lake, Syria.

Pocock.

Ibidem,

p.

720.

Perevyazka
tury,
it

is

now

an artificial, bookish name, though in the 18th cenin

use in the Ukraine (Pallas, 1811). Perhaps it is connected with the striking white transverse band ("bandage") across the facial part. In the steppes of the northern Caucasus, in particular along the Terek (V.G. Heptner) and in Kuban' (Dinnik, 1914), the name "berguznaya" existed widely. The name given by Gueldenstaedt itself represents simply a Latinized Russian word. Long ago (18th century) among furriers and sometimes also now,

was apparently

name "polecat-perevozchik" or simply "perevozchik" is used. This name is It may itself represent a distortion of one of the old Ukrainian folk names "perevyznik" (Pallas, 1811).
the

without meaning.

*In Russian original, rendered as "Tiveriadsk"

Sci. Ed.

1181
1936. Vormela peregusna ornata. Pocock. Ibidem, p. 721. Near

Lake Baikal

Nomen

delendum! (V.H.).

1948. Vormela peregusna pallidior. Stroganov. Tr. Zool.

SSSR, 7, p. 129. Semirech'e, former Kopal'sk 1948. Vormela peregusna obscura. Stroganov. Ibidem, Vakhsh valley, Tadzhikistan (V.H.).
Diagnosis

Inst.

AN
131.

p.

Only species

in its

genus.

Description
In general appearance, the
cats.

marbled polecat
its

is

very similar to pole-

As

in

them,

it

often

moves

in leaps, usually arching the back.

With
voice

this,

however, several of

ecological particularities are

noticeably different from polecats: instead of a sharp chirp, the

is

threatening

peculiar growl or grumble, and adopts a

special defensive pose, etc. (see below).

Head small and narrow, muzzle obviously

very large

blunt, ears relatively

tall

and broad

at base,

sharply delineated due to their

784

Fig.

277. Marbled polecat Vormela peregusna Giild. Sketch by A.N. Komarov.


1182

785

Fig.

278. Marbled polecat Vormela peregusna Giild.

Badkhyz preserve, southern

Turkmeniya.

May

1962.

Photograph by Yu.K. Gorelov.

coloration. Tail long, constituting about half of

body length or

slightly more, covered with long hairs, very fluffy.

tively

Winter fur quite short and not very close-fitting, with relaweakly developed underfur and dense guard hairs. Fur, not

counting very long hairs on

tail, is even, and sharp contrast between underfur and protruding guard hairs, as in polecats, is absent in marbled polecat. Winter fur quite soft. Summer fur somewhat shorter, sparser and coarser than winter [fur]*, but in

general, differences in characteristics between winter and

fur relatively small, and

summer

much

less than in

northern forms of

subfamily, even, apparently, less than in steppe polecat.

Anal glands present. Teats 5 pairs (4 abdominal, 1 thoracic; Ognev, 1935). Color of the marbled polecat is to the highest degree unique it is very variegated and itself consists of a combination of black, yellow and white areas and spots. It may be interpreted as a combination of facial pattern of the "mask" type (apparently, the extreme degree of development in the family) and a saddle-patch**
*ln Russian original, reads "summer," a lapsus

Sci. Ed.

**In Russian, chebrak, from Turkic shabrak, or saddle blanket

Sci. Ed.

1183
pattern. In general,

it

may be

considered that

it

corresponds with that

is

type of color which, within the family Mustelidae,

developed

in

honey-badgers, badgers and African polecats of the genera Poecilogale,

Leptonyx* and Poecilictis, but purely morphologically,

itself

appar-

ently represent an extreme degree of development and complication.

Color of the marbled polecat

cally.

is

not only complicated, but

entirely variable individually and, to a lesser degree, geographiIn

main features
(after A. A.

it

may be

represented by the following

scheme

Byalynitskii-Birula from Ognev, 1935, with

modifications and additions) according to the four main color fields

(Fig. 282)^.

Color of whole venter of body from throat to base of

tints.

the

tail,

including lower part of sides and legs, shining black color

with light brownish

Bases of hairs on belly are grayish-

brownish. Sometimes

in

middle of posterior part of belly, small

light-rusty dots occur, or several light spots are found.

Head
786 pure

color very clear and, contrasting, consisting of black and

lips,

white parts. Upper and lower

and chin

back corners of mouth "peristomal ring" pure white. Quite wide black
to
it

stripe

passes through eyes in form of transverse band. Descending to

sides of head and reaching corners of mouth,

unites with black

throat area.

Behind

this

black band, a quite broad pure white

transverse stripe runs between eyes and ears

"frontal band".
it

On

reaching lateral head surface of the head,

posteriorly, passing

slopes

below ear and ending on sides

somewhat of ear. Remain-

ing part of head behind white frontal belt, including occiput, black.

Ears located on this black area. At the base and posteriorly they
are black; edges support broad pure white margins of

somewhat

longer hairs. These bright ear margins

make

already large ears

look even larger. Bare

tip

of nose black.

Upper

side of neck dark, brownish-black with light pattern

consisting of three fields

two laterally which are wide and a widens anteriorly. Width of lateral stripe and degree of development moderately variable it is sometimes broken into several spots. In some cases, anterior parts of all three neck stripes themselves united in transverse direction, thus formnarrow middle one
that

ing another transverse,

more or

less wide,

white stripe

at

base of

*A synonym of Amblonyx, the African clawless otter, which lacks black and white coloration. Leptonyx is a lapsus, Ictonyx clearly being intended Sci.

Ed.

^Description given mainly after southern Russian form and schematically

because of
variation".

its

great variability. For additions, see below, and section on "Geographic

1184

nape corresponding as

it

were, to frontal band. Together with

lateral stripes (in these cases, the

middle one

is

poorly defined),

forms an angular bracket-like figure opening posteriorly

it

"collar".

Color of neck stripes white

somewhat yellowish

in posterior part,

and main tone of

this part of

neck brown or rusty-brown.

tail

Whole

dorsal side from shoulder region to base of

It

or

almost base, occupied by variegated saddle-patch.

spreads

ventrally and laterally, especially in posterior half of body,

it

where

also covers upper part of thigh. In anterior part, saddle-patch

bordered laterally by two usually quite broad white or yellow,

somewhat curved scapular

stripes.

They extend backward ap-

proximately to posterior border of thorax or somewhat farther.

Directly, or through series of spots, they pass to analogous stripes

bounding saddle-patch in its posterior part thigh stripes. They have same color, but on the whole are much less defined than scapular ones. At base of tail, along flanks, lie two oblong, light anal* spots. They are quite variable in size; sometimes each of them is divided into two, or there is a connection between them.

The
basically

actual saddle-patch itself represents a large area of

brown

tone.

Throughout

it

are scattered coarse yellow,

sometimes very light spots of irregular form. Often these spots flow together in longitudinal direction and form small, quite variable stripes. Nearer to the edges of the saddle-patch and along the

very edge, the tendency towards fusion of spots and formation of

stripes is stronger,

and here they often completely fuse with the

thigh stripes, from

whence the

latter loses its definition.

The basic tone of

the saddle-patch

may

also be yellow and in

this case, relatively small,

color are scattered over

it.

more or less rounded spots of brown They also sometimes display a tendency
is

towards fusion into longitudinal, oblique, and even transverse

groups. The anterior part of the saddle-patch

commonly more
is

densely covered with spots; in

its

posterior portion they are smaller,

field

more sparsely
paler,

scattered,

and the yellow

of the saddle-patch

on the whole, more clearly defined and more

striking to the eye.

Both described types of saddle-patch themselves represent in saddle-patches with light main field, this field is formed by a strong increase in area and number of fused yellow spots, and the brownish spots on it are, apparently,
negatives of one another

*In the Russian original, "analnykh (poroshitsevykh)"; both words appear to have the same meaning Sci. Ed.


1185
the remains of the
first type.

main dark background of saddle-patches of the

is to

On
all

the whole, saddle-patch color

the highest degree

the color type

variable, both in general features

in

and one

details.

From

which

basic elements of the saddle-patch are expressed, are

side, as far as a large, light,

787 derived the origins of all forms;

area with insignificant dark spotting, and on the other hand

a dark

up

to

dorsum with a

relatively very small

number of

light spots.

In the first case, thigh stripes completely disappear, while scapulars

are poorly developed, and in the second, scapular stripes are very

well expressed and thigh stripes are

commonly

sufficiently

marked

they are continuous or broken into individual spots. Scapu-

lar stripes are usually well differentiated in all cases.

Its middle segment with brownish highlights, and the tip is bright blackish-brown. Color of individual hairs at their base and proximal part is pale-rusty-whitish, and distal part is rusty-brownish and bright. Base of hairs on middle part of tail whitish, passing farther to a brownish-rusty and in part brown band, and hair tip white or slightly yellow. Lower surface of tail
tail is

Base of

rusty-brownish or brown in color.

a pale-straw

the longest part

is

at

base region darker than upper, usually bright blackish-brown

feet

and on remaining part, as on upper side. Vibrissae and hairs covering soles of
pads not covered with hairs
in

dark-brown, foot
light,

winter.

Claws

quite long,

weakly curved.

The above-described deviations in color do not comprise the whole range of its variability. Individual parts of pattern on head vary in color and form. The [white] peristomal ring occupies the chin to a variable extent (except for lips, it may be entirely black); the white frontal band may descend on the sides of the head and neck to a variable degree sometimes ending below the ear, sometimes passing farther onto the throat, and in some cases, being interrupted on the forehead by black, in others uniting under the eye with the peristomal ring, etc. Sometimes, light spots appear on the middle of the abdomen as if uniting the saddle-patch edges from opposite sides. Finally, the intensity of color varies in both dark black and brown and light areas yellow and pure white

color.

All the changes mentioned are mainly of an individual character;

however,

in

some

characters geographic variation also reveals

itself.

This concerns general tone of saddle-patch and the relative

1186

degree of development of dark and light tones of its basic background, sharpness of pattern of scapular and thigh stripes, general intensity of ventral black tone, appearance of light spots

on abdomen,

etc.

Sharp mutational changes

in color

have not been

described in the marbled polecat.

Sexual dimorphism in color is absent. Color of young differs from adult color, the elderly being more intense and darker in basic brown color of saddle-patch, its spotting being lighter with more noticeable development of yellow tone. The tail is less fluffy.
Seasonal variation in color is not expressed or is weak. For characteristics of skull, see above in characteristics of the
genus.

The
is

structure of the os penis

is in

the

main similar

to majority
Its

of species in genus Mustela in particular, to the polecats.

laterally

base

compressed, and the

flattened.

distal third is

bent upward hook-

like.

The

dorsal surface of the basal half carries a rib-shaped elis

evation, and the end

The very

tip itself

denticle slightly bent to the right. There

is
it

forms a rounded no groove on the lower

is

surface of the basal part, but terminally

well-developed.
to great

Dimensions of the marbled polecat are not subject

not been noted.

individual variation; in particular, especially large individuals have

269-352 mm, of females, 290-345 18-205 mm, of females, 159-184 mm; length of hind foot of males is 39-50 mm, of females, 38-45 mm; ear length of males is 20.5-31.0 mm; of females 24-27 mm (from approximately 90 individuals, after Ognev, 1935; Pocock, 1936; Stroganov, 1948, 1962; Chernyshev, 1958 and material of the

Body
tail

length of males

is

mm;

length of males

is 1

Z[oological]

M[useum

of]

M[oscow]

U[niversity]).

Condylobasal length of male skull is 50.6-62.0 mm, of fe789 males, 49.0-56.3 mm; zygomatic width of males is 29.9-39.0 mm, of females, 30.0-34.9 mm; interorbital width of males is 13.1-19.0

mm,
32.6

of females, 13.1-16.0

mm;

mastoid width of males

is

26.1-

mm,

of females, 27.2-30.2
is

mm.

Os

penis length
is

36.9-39.2

mm (3
(V.H.).

individuals;

Ognev, 1935).

General weight

370-715

gm

1187

788

Fig. 279. Skull of

marbled polecat, Vormela peregusna

Giild.

1188

Systematic Position

Only species

in the genus.

Geographic Distribution

Found

in steppes, semideserts

and deserts of southeastern Europe,

Near East, Middle Asia and part of Central Asia.

Geographic Range
in the Soviet

Union

The range
half.

(reconstructed) constitutes a considerable part of the

itself represents the

range of the species and

north of

its

western

boundary of the range begins at Belovezhsk Forest (approximately 53" N. lat. and 24 E, long.). Thence, it is directed to the southeast, to Kiev, and then rises again towards the north, reaching the Maloarkhangel'sk region (at the source of the Oka 52 20'), is directed eastward to the Voronezh
In the west, the northern

region (Voronezh preserve) and thence, in an imprecise way proceeds to the Volga to Syzran' or a little south of Syzran'. Beyond

Volga the border, apparently, rises north of 52 [N. directs itself to Orenburg on the Ural river.
the

lat.]

and

to the foot of the

range in the west extended, apparently, Carpathian mountains but in these mountains, the marbled polecat was evidently not present. It was also not recorded in the Trans-Carpathians. The range probably extended into

South of

this line, the

Romania

polecat existed

extreme lower Danube. Apparently, the marbled over Moldavia. Farther to the east, the range of the marbled polecat everywhere reaches the shores of the Black and Azov seas, occupies the Crimean steppes (marbled polecat is absent in montane Crimea, but was noted at Balaklava) and extends to the shore of the Caspian Sea. In the Caucasus, the marbled
in the
all

from Makhachkala, reaching the foot of the mountains (Nal'chik, Vladikavkaz). In places, it goes up into the a tributary of Kuban'), foothills somewhat (Psebai on the Belaya but in wooded montane regions, it goes no higher than 600-700 m above sea level. It is found along the entire Caspian shoreline to
polecat
the
is

distributed over the entire Cis-Caucasus steppes

the

mouth of

Kuban'

to

Derbent, and farther south, to Baku.

1189
In the Trans-Caucasus, the marbled polecat inhabits the plains

Kura and Araks basins, passing along the Kura it is met with along the valley of the Araks and the montane steppe region and in semideserts north of it,
and
foothills in the
to Tbilisi. In

Armenia,

west of [Lake] Sevan) to Kirovakan region and Leninakan (latitude 40 15' N. lat.) and somewhat north of it to Gukasyan region (41 00' N.
proceeding into the western part of the country
(to the

lat.).

In this part of the Trans-Caucasus, the marbled polecat

is

found

at a height

of 1700 m.

It is

encountered in the Lenkoran depression

and

in unforested places of

Talysh

at

heights up to 2000 m.

Beyond

the Urals, the northern border of the range

from

Orenburg and the Ural river passes southeastwards to the upper

Emba, and then swings

to the east and, occupying the Pri-Aral'sk Karakum', is directed to the Ulutau mountains, to the upper Sarysu and Semipalatinsk. Thence, it enters into the Cis-Altai steppes and goes on to Rubtsovsk and Biisk. In the Biisk region, it swings sharply around to the southwest (the eastern end of the range forms

a sharp extension) goes along the Altai foothills including Ust'-

Kamenogorsk region and Katon-Karagai, and

sion,

the Zaisan depres-

and here exits beyond the borders of our country to Dzhungaria

the south of the above-mentioned line, in the Asiatic part

[China].

To
plains.

of the country, the range occupies the whole of the Middle Asian

The southern border

either goes

beyond the borders of our

country, or passes along the foot or foothills of the southern moun791

Thus, the range does not include Tarbagatai and Dzhungarsk Alatau, but passes between them, as well as along the Il'i valley and Il'iisk depression to China. Farther west, the southtain systems.

ern border skirts the Tien-Shan system; apparently, the Fergana

Valley together with Karatau are included in the range, and

skirts

it

around the Pamiro-Alaisk montane country. It crosses Zeravshan valley near Pendzhikent and passes around the Baisunsk mountains or occupies their southern foothills and Kugitangtau. To the east, the marbled polecat is encountered in the Surkhandar'ya valley, at least up to Denau and Saryassiya, in the Babatag (between Surkhandar'ya and Kafirnigan), along the lower Kafirnigan (apparently, not reaching Dushanbe); along the Vakhsh, it penetrates as high as Kurgun-Tyube, and along the Pyandzh it apparently reaches the mouth of the Kyzylsu, and perhaps to Kulyab.
Apparently, the animal
is

more widely

distributed in southern

1191

Tadzhikistan along river vallies and low dry mountains than has

been shown. Besides being an undoubted inhabitant of the plains, however, the marbled polecat penetrates along valleys high into the mountains in some places (Tien Shan) and is even recorded at heights up to 3000-32000 m.
exits

the west of the Amu-Dar'ya, the range of the marbled polecat beyond state borders everywhere. It is, however, absent in the Kopet-Dag, although in the lower western parts of the range, it

To

penetrates into the mountains, notably along the foothills and valleys.

the inhabitation of the marbled polecat Trans-Urals to 54 50' N. lat. (Kundrava southeast of Chelyabinsk not 55 30' as written by Ognev, 1935) is a mistake and
in the

The assumption about

is,

apparently, based on an imported skin.

existence around Baikal,

scribed,

whence

The affirmation of its separate form was even dewell-founded

and
is

in "Siberia" (Pocock, 1936) is not

this

species

absent everywhere east of the Cis-Altai steppes in our

country.
In the last 100-200 years, the range of the marbled polecat has been contracting significantly. This decrease started from the west. Thus, information about the occurrence of the marbled polecat in Belovezha reflects the boundary in the 18th and 19th centuries; in Volyn' and Podolia" it existed in the first half of the last century; in Moldavia ("Bessarabia") it disappeared about 100 years ago.

Near Odessa, the animal existed even at the end of the first half of the last century, but by the 70' s, it had already disappeared and later appeared occasionaly and very rarely (Fig. 281). In the west, beyond the Dnepr, this animal is now absent and almost everywhere it has even been completely forgotton. At the beginning of our century (before 1914), the marbled polecat was only recorded twice on the right bank of the Dnepr near Kiev and near Odessa. These were, obviously, transgressions, although it

cannot be excluded that near Odessa,

it could have appeared from Romania. From there (from Dobrudzh) transgressions to Izmail'sk district are not excluded even now. The border has shifted very strongly eastward, even east from the Dnepr. Occurrences near Pavlograd and around Zaporozh'e have to do with the beginning of our century (to 1914). In vast expanses, though not everywhere, between the Dnepr and the Ural,

the northern border of the range has also shifted to the south.

1192
Thus,
at the

beginning of the current century

still

in

Orlovsk

district,

the marbled polecat

it

existed, although

it

was

rare, but in 1925,

later, the

had already completely disappeared. At that time or somewhat marbled polecat also disappeared from Kursk district. In the 50' s of the present century the western boundary of the marbled polecat's range was represented a line of complicated outline (details insufficiently known), going at first from Voronezh

(Voronezh preserve) southwest to Kupyansk or to a place between them and Khar'kov, and thence to Krasnograd already the right bank of the Donets. Here the border swings sharply southeast and passes through Artemovsk to Lyugansk. Somewhere on the right bank of the lower Donets, the border swings back sharply to the southwest and goes straight west some distance from the shore of to the the Sea of Azov to Osipenko city or a little westward

meridian of Melitopol.^
792

In the interfluve

border, begins near Voronezh, descends

then, passing slightly north of 50 N.

between the Dnepr and Volga, the northern somewhat southward, and
lat.

directs itself eastward to

Khoper. Thence,

it

crosses over the Volga at an

unknown

point, to

the middle course of the Uzenei on the interfluve of the

Volga and

Ural [river] (Saratov

district).

Individual occurrences of the marbled polecat were registered,

already long ago, both west and north outside the outlined area of
present distribution
less

these

are Poltava (perhaps even a

in

more or
100-200 mention

permanent occurrence), and Pugachev

Trans-Volzh'e.

On

the whole, the border of the range for the last

years receded very strongly

the right

from north to south (not to bank of the Dnepr) for 350-600 km, and from west to east for 700-1000 km. The latest shift of the range led to that this "rounded pontoon" ring which the primordial range formed in its western part was shown to be broken, and the union between populations of extreme southeastern Europe (Romania, Bulgaria) and the southeastern part of our country (Don and Volga steppes) was already absent. Reduction of the range in the south and east
in the European part of the country continues and apparently, the complete disappearance of the marbled polecat, at least in the expanse between Dnepr and Don, is not far off. At present, the

^Sokur (1960) demonstrated

region); however, this place

its

occurrence also in Nikolaevsk

in the

district

(Vladimirsk

was not shown

map and no

details are reported.

1193

marbled polecat is already only sporadically distributed, rare or even very rare everywhere west of the Don; in many regions it appears occasionally and is obviously on its way to complete
disappearance.

The reason

for the

marbled polecat's disappearance

it is

lies in the

ploughing of the virgin steppes with which

associated, and in

the complete agricultural utilization of the steppe and forest-steppe,

and

in particular, in the

area of their normal and

gins east of the

Don

(at

disappearance of ground squirrels. The more or less continuous distribution beKamensk, Shakt and others), and in the

Cis-Caucasus.'*
It is

very interesting that reduction of the range of the marbled

polecat in Europe proceeds in parallel the expansion of the steppe

polecat's range. Replacement of a species associated with the
virgin steppes
its

"a

fugitive of culture"

by

another carnivore of
but revealing a

lineage ecologically analogous in

many ways,

tendency towards being a "companion of culture". They secure for themselves subsequent prosperity where the marbled polecat
does not
cies
persist.

No

competitive relationships between these speof sympatry.

was observed

in sufficiently extensive areas

In the east, reduction of the range remains almost unobserved.

Changes recorded there belong, apparently,

to the

category of

negligible natural fluctuations in the border of the range.

Fossil remains of the marbled polecat have been found only

within the limits of

its

recent range.

Geographic Range outside the Soviet Union

Europe the range includes Dobrudzh in Romania (in remaining Romania, only transient in Krainovo district near Danube), all of Bulgaria, eastern and southern parts of Yugoslavia (Serbia, Kosovo and Mefodia, northern Macedonia, southern Adriatic coast; in Slovenia, transient in extreme northwest of country, European
In

part of

"Range according to data of Brinken, 1829; Kessler, 1850; Eversmann, 1850; Greve, 1894; Miller, 1912; Dinnik, 1914; Satunin, 1915; Brauner, 1923, Averin, 1928; Migulin, 1929, 1938; Calinesscu, 1930, 1931; Nezabitovskii, 1934; Ognev,
1935; Flerov, 1935; Pocock, 1936; Ptushenko, 1936; Charleman', 1937; Vereshchagin, 1947, 1959; Kuznetsov, 1948, 1952; Stroganov, 1948, 1962; Komeev, 1952; Sludskii,

1953; Dal', 1954; Heptner, 1956; Barabash-Nikiforov, 1957; Chernyshev, 1958; Afanas'ev, 1960; Sokur, 1960; Ishunin, 1961 and other sources and unpublished material of N.I. Larina and V.G. Heptner.

1194
part of

Turkey and easternmost part of Greece, west approxi-

mately to Salonika (Calinescu, 1930, 1931; Brink, 1958; Markov,

1959; Dulic and Tortric, 1960).
In Asia, the range occupies the Mongolian Republic north to 47 (southern foothills of Khangai, Undurkhan) and to 49 (west-

793 ern great lakes region). Inner Mongolia, including Ordos^, Afghanistan, Baluchistan, Iran, Iraq, Syria, Palestine

details of

its

distribution here are

and Asia Minor (the unknown)*. Probably, the

is

range goes on to Kashgaria, but positive information about this

absent (V.H.).

794

Geographic Variation
Geographic variation of the marbled polecat
In
all,

is still

poorly studied.

about 12 geographical races are known, of which more than

half belong to our territory. This

ated, for there are not
try.

number

is,

apparently, exagger-

more than

three real subspecies in our coun-

coloration.

The fundamental characteristics of all forms are based on At the same time, as shown above coloration of the
is

marbled polecat

subjected to very strong individual variation. In

several cases, general dimensions or claw length were given as

characters, and craniology not given at
all.

Great variation of color,

ple,

commonly based on
is

limited material,
(for

was presented by several authors, especially older ones

K.A. Satunin) but also recent,

exam-

clearly in error. All these are

responsible for contradictions, both in interpretation of various

forms, and also in descriptions of their color characters as well as

distribution. Thus, in the opinion of
in

one author (Stroganov, 1948), the nominal form the white band on the forehead is broken,
all

while

other authors describe

it

as continuous; in another case,

to

the distribution of the

form negans was found

it

be disjunct, and

between the two parts of

1956), etc.

lies the

range of another form (Novikov,

Based on considerable material (Z[oological] M[useum of] M[oscow] U[niversity]), a preliminary scheme is given below of the geographic races recorded in our country, and in part, in
'Shanxi but not Shaanxi as some authors write,
i.e.

inside the great northern

bend of the Huang He and not on the left bank of the river west of it. *But see Harrison and Bates, 1991. The Mammals of Arabia, 2nd
Zoological Museum.

ed. Harrison

1195

793

Fig. 281.

Some

previous and present borders of range of marbled polecat, Vormela

Giild, in the

peregusna
1

European part of USSR. V.G. Heptner:

western borders of range (western more or less beginning of 60' s) northern and western borders; Separate distant transgressions in in last decade; 4

approximate); 2
3

Reconstructed northern and The recent Separate points of occurrence

(at

decade (Poltava, Pugachev); 5 ning of 20th century (before 1914).

last

Some

peripheral places of occurrence in beginin

Map depicts formation of the break pontoon" ring of original range.

"rounded

1196
adjacent ones.

It

was ascertained
lies

that the

main feature of geo-

graphic variation of color

first

not in detailed color difference, but

of

all,

in its

general tone, mainly on the dorsal body, and in

the ratio of darker to lighter colors (density of spots, their general

area) in the saddle-patch.

Since statements of some authors (Stroganov, 1962) on differences

in size

between races have not been confirmed, dimensions

V. p.

of the separate forms are not given.

1.

South Russian marbled polecat,

peregusna Giielden-

staedt, 1770 (syn. sarmatica, intermedia, euxina).

General color dark. Basic prevailing color of dorsal saddle

dark rusty-brown or brown and sometimes dark- or blackish-brown. Light spots on
it

relatively small

and few, bright brownish-yellow,

yellow-straw or whitish-yellow color. Three longitudinal occipital

stripes,

and

lateral separated

from dorsal and not fused anteriorly

(no "collar" formed). Color of occipital stripes pale-yellowish, some-

times white. Scapular stripes well marked, thigh stripes often

however, they always bear dark spots. The white frontal band not divided medially, and does not unite with peristomal white ring or very rarely forms one. Abdominal spots absent, or in all 1-4 may occur and they are small and dull. In the south and southeastern European part of the country, Cis-Caucasian steppes and probably the Trans-Caucasus. Boundaries with respect to the next form unknown. Outside the USSR in Romania and the Balkans. The systematic position of the marbled polecat of the TransCaucasus has not been studied in the manner required. They have been related (Ognev, 1935) to the population inhabiting the eastintermedia') on the basis of very little mateern Cis-Caucasus rial. It is possible that they belong to syriaca a form which in
distinct;

general
2.

is

very

little

known.
V. p.

Transcaspian marbled polecat,

koshevnikovi Satunin,

1910 (syn. alpherakyi, tedschenica, obscura). General color dark. Basic prevailing color of dorsal saddle 795 dark-brown or even blackish-brown similarly dark or darker than the nominal form and with less red tinge. Light spots few, for the

most

part light-yellow or white in color. Occipital stripes united

by a broad transverse white stripe (forming well-defined "collar"). Color of lateral occipital stripes usually yellowish. Scapular stripes very distinct; of thigh, weakly defined and bearing dark
anteriorly


1197

spots.

White

frontal

band not divided medially and has no connec-

tion with peristomal ring.

occur, weakly defined and few (1-3).

less than 10

Abdominal spots are absent or rarely Claws short the longest

mm.

eastwards to at least AmuIn the western part of Middle Asia Dar'ya and region of right tributaries of Pyandzh and upper AmuDar'ya (southern Tadzhikistan). Distribution eastward beyond Amu-Dar'ya and boundary with next form not clear apparently, however, does not pass to east of Karatau [range] (see below). Border with nominal race also unclear. Outside the USSR Iran, Afghanistan, West Pakistan

(Baluchistan).

The Transcaspian marbled polecat represents

race, close to the

a well-defined

nominal form.

It is

very similar to the latter in

dorsal color, but dark tone often even darker and therefore fur

color appears
ter,

more

contrasting. Except for this indistinct charac-

both forms are well differentiated from each other by the

presence of a "collar" in the Transcaspian form.

The dark form described from southern Tadzhikistan (obscura)

has in general a completely similar type and intensity of color.
Characteristics of head coloration mentioned by the authors of the
796 original description are not constant,

and do not have essential The author's suggestion concerning its small dimensions not confirmed (Chernyshev, 1956). Although the
diagnostic significance.

author considers this form "one of the most differentiated subspecies" (Stroganov, 1948), there
it

is

insufficient basis to separate

(valleys of

from the Transcaspian V. p. koshevnikovi. Moreover, its range Vakhsh and Pyandzh) is perhaps very small. Such small

ranges are not characteristic of races of Palearctic carnivores.

The forms peregusna and koshevnikovi

and, apparently, to
its

as a pair of closely

related dark forms, contrast sharply with the very light pallidior
pair

member
in

negans.

The presence of dark

ters

races which

shows no

essential charac-

of "degradation"

of color

Turkmenia and southern

Tadzhikistan and, apparently, in Afghanistan

defined desert climate and in general, desert conditions

countries with

clearly
is,

in

some

respects paradoxical. In a series obtained from Turkmenia,

one may find skins having purer, more saturated black tones, generally with more contrast (brightness) than among the nominal race. It is notable that such races occur even in the middle part of

1198

<

795

Schematic drawings of color of several forms of marbled polecat, (from Ognev, 1936, with modifications). Upper row extreme types of color variation of South Russian marbled polecat V. p. peregusna Giild (on right specimen from Cis-Caucasus); lower row Transcaspian marbled polecat V. p. koshevnikovi Sat. (on left) and Semirech'e marbled polecat, V. p. pallidior Strog. {"negans"). There are substantial differences in the form of the lateral occipital and finely dotted stripes as well as general development of black and light colors on
Fig. 282.

dorsal saddle.

1199

796

Fig. 283.

Marbled polecat surveys its surroundings before coming out of its burrow, Kyzyl-Dzhar village in Badkhyz preserve, southern Turkmeniya. May 1962. Photo graph by A. A. Sludskii.

the

U[niversity]).

Kara-kum (Uch-Adzhi; Z[oological] M[useum of] M[oscow] The very light color of the Semirech'e and Central

Asiatic animals clearly contrast with this form. There are, therethe dark European, fore, two groups of race of marbled polecat Near Asiatic (Afghanistan, Iran) and western Turkestanian races, and the light Semirech'e (eastern Middle Asiatic) and Central Asiatic races. Such a division of race groups reveals certain analogous features known in some other carnivores (weasels, badgers,

bears).

The name adopted here

is

for the Transcaspian

form nas an

in-

disputable priority over alpherakyi* Birula, under which this form

usually known, although both of them appeared in the

same

year.
3.

Semirech'e marbled polecat,

light.

V. p.

pallidior Stroganov, 1948

(the
797

name negans was

General color

applied to this form).

Prevailing color of dorsal saddle light,

changing from ocherous-yellow to almost white, often straw yellow. Over it, are scattered sparse tawny, brown or rusty-brown
"In

Russian original misspelled alhperakyi

Sci. Ed.

1200
spots.

Occipital

spots

in

its

anterior

part

connected by a

transverse white stripe ("collar" well developed). Color of lateral

occipital stripes whitish. Scapular stripes not sharply defined, but

due to generally light color of dorsal saddle, are well marked. Thigh stripes undefined or almost undefined and strongly covered with spots. The white frontal band often divided by narrow (up to 1 cm) medial black stripe, but has no connection with white peristomal white. Light spots are encountered fairly often on abdomen and are usually larger. Claws quite long reaching 13 mm. on

middle digits of fore

Altai steppes and

foot.

In the region to east of the Karatau range, Semirech'e, Cis-

Outside the

USSR in

montane steppes of southern Altai (Chuisk). Dzhungariya, and Mongolian Republic,

apparently except in southeastern or eastern parts.

well-distinguished form, obviously contrasting with both

It is

previously described forms.

species, or in any case of

possibly the lightest form of the

the group of light races (see

members of
it

above). Based on

its

color type,

represents a sort of negative of

the races of the dark group

the

ratio of

dark to light areas on the

dorsal

body and saddle are reversed.

comparison
to several other races
is

Despite the obviousness of characters of the Semirech'e population in the sense of

and

distri-

butions

this,

of the

many

forms,

the least

known and
V. p.

unclear.

The Semirech'e marbled polecat was previously (Ognev, 1935)

considered as transitional from the Transcaspian
koshevnikovi

Calpherakyr)

to the Central Asiatic (Ordos) V. p.

its
is

allocated directly to this race. In any event,

negans or was differences from the

completely correct.

Transcaspian form were emphasized, which

clearer delimitation between the forms pallidior and negans

than that followed (Stroganov,

1948)

is

required.

On

the other

hand, the limits of the distribution of this form to the west are

unknown. The suggestion

that

it

is

distributed "from the eastern

slopes of Karatau. ..southward, apparently, to Turkestan and the

Alaisk (in the text, in error as Altaisk; V.H.) range" (Stroganov,

1948, 1962),
the area
is

not understandable. In general, the race inhabiting

between the Amu-Dar'ya in the west, Karatau in the east, and Turkestan and the southwestern spurs of the Gissar range in the south is not clarified. In any case, considering the Semirech'e pallidior, there is no basis for relating animals from Uzbekistan and Tadzhikistan to the form negans as is sometimes done (Novikov,

1201
1956). This view was, apparently based on misunderstanding since

such a gap in the range of a race

is

improbable.

Note. Pocock (1936) described a separate form of marbled

polecat V. p. ornata from "the vicinity of Lake Baikal". Judging by
the photograph, this form
is

of moderately dark color with a well

is

developed "collar";
the marbled polecat

i.e.,

an eastern type. However, as

known,
(if

is

absent not only in the vicinity of Baikal

one may speak of proximity to a lake extending for about 1000 km from north to south), but also in a great area around it. The author had 2 more specimens from "Siberia", and he proposes "Siberia"
as the region of distribution of this race.

To

establish that

it

particularly represents the

and

this

name

is

form ornata is not in any way possible best excluded in the category of nomen delendum.

Beyond
Miller,

the borders of the

USSR,

the following forms are

V. p.

indicated (not counting euxina, see above, and ornata):

1910

Ordos,

negans

Inner Mongolia, Shaanxi and

some other
p.

parts of Inner Asia, not accurately clarified,

and possibly also eastern

V.

or southeastern parts of the Mongolian Republic;

syriaca

Pocock, 1936
to
798

Syria and adjacent

territories (Palestine?)

eastward

western Iraq.
Note.
V. p.

chinensis Stroganov, 1962 described from a speci-

men

labeled "China" and earlier assigned to negans (Ognev, 1935)

status.

does not deserve separate

origin of the type)
is

The nature of area inhabited (and

of the lower course of

set forth as "basin

Hwang Ho". None Hwang Ho

absent.
if that

of the arguments in use leads to such an interpart of the river

pretation of the label, not to mention that in the lower course of

below the great northern bend

is

can be so considered, the marbled polecat

entirely, apparently,

At the same time, it was shown (Bannikov, 1954) that, judging by the route of the collector (Pyasetskii), this marbled polecat was caught at the southern border of the Mongolian Republic, i.e. in the region inhabited by the form negans and not far from its type locality (V.H.).

1202

Biology
is encounEuropean part of the country they are few in number; it is, with few exceptions, a rare animal. In the Don and Kuban steppes, it was comparatively common in the 20' s. In the CisCaucasian steppes, the marbled polecat was most common in the eastern part of the steppes (Kizlyarsk steppe) where it is drier.

Population. Almost everywhere the marbled polecat

tered in the

There

it is

most often met with

is

at the

present time (V.G. Heptner).

in

The marbled polecat

now common

Kazakhstan,

in

the

Muyunkum
(Sludskii,

sands, southern Pribalkhash'e, and along the Syr-Dar'ya

1953).

The marbled polecat

its

is

most

common

in

Turkmenia, where
great gerbil

distribution

is

closely connected with the

it is

[Rhombomys opimus] and

Its

encountered formerly
it

in

great numbers, in the flat part of the country

was

the

most

common

small carnivore.

population significantly exceeds the

weasel population (V.G. Heptner).

Habitat.

The marbled polecat

is

an animal of open
It

unwooded
met with

expanses

dry
range.

steppes, semideserts and deserts.

is

chiefly in places rich in large desert and steppe rodents

gerbils,

ground
duces
the

squirrels

and
It is

in part jerboas, etc.

The ploughing of

virgin

steppes and fallow lands pushes out the marbled polecat and reits

encountered

in the

southern grass steppes of

chernozem zone, does not avoid thickets of blackthorn, buckthorn and other shrubs in steppe shelterbelts. It infrequently occupies afforested river valleys and the borders of forest massifs of
799 the forest-steppe.
It

is

encountered occasionally among gardens,

but in dry hilly steppes,
localities of desert
it

melon

fields or

even

in settlements;
it

is

rare. In

Middle Asia,

mainly inhabits
it

and

does not avoid river valleys and sections cut by irrigation ditches and sometimes overgrown with a
semidesert character, but

border of tamarisk.

It is

most

common

in stabilized slightly hilly

sands, sprinkled with black saxaul, winterfat, pea-shrubs [Caragana]

and milk-vetches [Astragallus] alternating with solanchak with scattered saxaul woods. In sagebrush-grass semideserts, serozem
sagebrush deserts, clayey steppes and sheep's fescue-feathergrass foothill steppes, it is met with rarely (Sludskii, 1953). The marbled
polecat prefers the plains and for the most part does not go higher

than the foothills.

It

inhabits woodless mountains covered with

1203

798

Fig. 284.

Marbled polecat running in grass. Badkhyz preserve, southern Turkmenia. May 1962. Photograph by Yu.K. Gorelov.

montane steppes. In Talysh, it is rarely found more than 2000 m above sea level and in Tien Shan up to 3000 m. Food. Information on foods of the marbled polecat is scarce. Apparently, of greatest significance in Middle Asia are the various species of gerbils, especially the great gerbil with which it is always closely associated, and the long-clawed ground squirrel [Spermophilopsis leptodactylus]. The marbled polecat feeds on other ground squirrels [Spermophilus], jerboas, hamsters and various

mouse-like rodents. In captivity

it

eats small animals, birds, lizards

and eggs. Cases of attacks on domestic fowl are unknown.

Home

range. Information
shelters.

is

absent.
for the

Burrows and

The marbled polecat

in

most

part

inhabits burrows of various steppe rodents, only widening and

deepening them. In Turkmenia, and generally

Middle Asia, the

(towns) of great
this

marbled polecat
gerbils, not only

lives

permanently

in colonies

because of the close trophic connection with

its

rodent but also in connection with

use of the complicated

underground structure as a diurnal shelter in case of danger and during reproduction. It sometimes uses burrows of long-clawed

1204

799

Fig. 285.

Marbled polecat

in

preserve, southern Turkmenia.

colony of great gerbil, Rhombomys opiinus. Badkhyz May 1962. Photograph by A. A. Sludskii.

ground

squirrels, but this

its

is

an exception. The marbled polecat

as a rule (V.G. Heptner).

does not make

own burrows

Daily activity and behavior. The marbled polecat is mainly 800 active in twilight and early morning. It spends the day in burrows

and autumn, occurs in the daytime activity when the heat is still not too great, marbled polecat or morning hunt is sustained longer when sun is
(Sludskii, 1953). In

Turkmenia

in winter time, spring

very high.

As

a kind of exception, the animal appears

on the

sur-

face during the day in

summer time (V.G.

Heptner).

The marbled polecat hunts

for great gerbils under the ground.

The

predator, penetrating the subterranean passages of a town,

causes great disturbance

among

its

inhabitants

the gerbils run out

from the burrows, hided again in them, try to run across to neighboring colonies, etc. The hunting marbled polecat appears first in one burrow and then another, disappears underground again,
and again appears, etc. In open expanses, following, for instance, widely wandering midday gerbils [Meriones meridianus], the marbled polecat rarely catches them; mainly it catches them in

burrows (V.G. Heptner).

1205

rt
f
^fi^^^"

^)
'*<
>.

^?'^-'> f '^^N

^
800
Fig. 286.

Tracks of marbled polecat on wet sand

in a gallop

prints are placed in fours

hind

and scheme of

leaps. Foot

feet

do not

fall in

the tracks of fore feet.

Karakum

80

km

to north of

Ashkhabad, 25 January, 1948. Sketch by A.N. Formozov, about

2/3 of natural size.

characteristic behavior of the marbled polecat manifests

itself in the intimidation

pose which

it

displays in case of fear. In

a defensive pose, the

tail is

raised especially strongly

it

back

(like a squirrel).
its

At the same time,

its
it

rises

up over its up high on its legs,

jerks up

head and bares

teeth. In contrast to the "chirring"

of polecats, ermine and weasels, dog. The frightening effect

is

gives a peculiar growl like a

strengthened by the bright, vari-

fur. When running, the marbled polecat expands its tail, fluffing it out. Seasonal migrations and transgressions. Data are lacking.

egated black-and-white

Reproduction. In the northern Caucasus, estrus occurs in March (Rossikov, 1887; Satunin, 1915). In Pribalkhash'e (Kazakhstan), a
pregnant female was noted
in the first

days of February (Sludskii,

1953). Throughout March, females which already had given birth

1206

young were caught. In the northern Caucasus, parturition later. On 9 May, there was a litter of six still completely helpless young (Rossikov, 1887). In Kazakhstan, parturition occurs, apparently, in February-beginning of March (Sludskii, 1953). The 8- week period of pregnancy occurring in the literature probably does not agree with actuality. The number of young in a litter ranges from 3 to 8. The average number (from 13 cases) is
to

occurs

4.3.

Thus, the marbled polecat

Males,

is

less

fecund than the steppe

polecat.
it

seems, participate in raising the young. Spermatogen-

esis begins to

be extinguished

in April.
is

Growth, development and molt. Information

801

absent.

Enemies, diseases, parasites, mortality, competitors, and population dynamics. This carnivore has

steppe polecat

may be named

as

no dangerous enemies. The one of the most fundamental

competitors influencing the distribution and numbers of the marbled polecat. Other steppe carnivores hardly exhibit a noticeable
influence as competitors.

The marbled polecat is almost unsusceptible more susceptible to tularemia.

to plague,

and

Human
to the
in their

changes

in the virgin

steppes and their ploughing lead

displacement of the marbled polecat and to a sharp decrease

the

number of

numbers. Utilization is not large and has no effect the marbled polecat (P.Yu.).
Practical Significance

at all

on

The importance of
is

the marbled polecat as a fur-bearing animal

negligible. In the

USSR,
Its

slightly

more than one thousand skins

is

are taken annually.

fundamental significance

as a predator infections.

which destroys steppe rodents, vectors of dangerous

Its

role in destroying rodents harmful to agriculture

is

is

not great

since the marbled polecat

rare in agricultural regions,

and

it

avoids cultivated landscapes.

(P.Yu.).

On

the whole,

it

is

a useful animal

1207

Genus of Honeybadger or Ratel*

Genus Mellivora
1780. Mellivora. Storr. Prodr. Meth.
ratel

Storr, 1780

Mamm.,

p. 34. tab.

A. Viverra

Sparrmann = Viverra capensis Schreber'.

Dimensions large. Limbs plantigrade, forelimbs with very long, strong claws, of fossorial type. Webbed membrane between digits absent. Skull massive and broad, with voluminous elongated braincase. Facial portion short and broad, the palate wide, paroccipital and mastoid processes large, strongly projecting (mastoid width is approximately equal to zygomatic). Auditory bullae large and swollen. Auditory tube covered from above by anterior part of mastoid process and opens below it. In general appearance, the skull is
entirely similar to skulls of representatives of the subfamily of true

martens.
Dentition of cutting (sectorial) type, with strongly developed
carnassial teeth (above, fourth upper premolars; below, first
lars);

mo-

molar relatively very small and strongly elongated in transverse direction. On the whole, dental system is quite similar in its main features (structure and relative size of teeth, etc.) to
first

upper

that of species of the subfamily of true martens, Mustelinae, especially polecats.

The

essential difference

is

only in that the second

lower molar

is,

as a rule, absent.

Dental formula

I-C-P-M- = 32. **
3
1

13 13

Ear pinna reduced and represented by short dermal ridge. Subcaudal glandular "pocket" absent, anal orifice located in a deep pouch radially striated by thickened skin; anal glands large and
gives off secretion with a sharp odor.

Pelage coarse and sparse, coloration of dorsal saddle type.

Teats, 2 pairs.

*The Russian common names medoed and

eater" and "bald badger" respectively

lysie

barsuk are

literally

"honey-

Sci. Ed.

'The generic synonyms, since they are not used in the Russian literature, are not
given.

**In Russian original, erroneously given as I

3^1 C"P M = 32 Sci.

3
1

Ed,

1208

802

An omnivorous predator, mainly feeding on small vertebrate and invertebrate animals. Ecologically, it is an entirely flexible form, encountered chiefly in different types of flatlands, mainly open,
to
forests;

sandy deserts inclusively, but it avoids continuous moist tropical it is met with mountains. Its capacity to dig is well developed.

The range of the genus is vast and includes the major part of Africa and Near Asia, India and the western parts of Middle Asia (for more details, see below, species description).

30

eo

%V^.

1>

Moscow'

eod

^=<5..,

Fig. 287.
ratel or

Range of genus of honeybadgers, Mellivora

Storr. and species range of honeybadger, Mellivora capensis Schreb. Schematic. V.G. Heptner.

1209

Within the boundary of the genus, many species were described, especially at the end of the last century and beginning of

the current one, both

present time,

it

from Africa and also from Asia. At the was established (Pocock, 1941) that the genus

includes only one species, with several races. This viewpoint

may

be considered generally accepted. The independence of the genus Mellivora has never been considered in doubt. However, the attempts made to establish its systematic position and phylogenetic relationships with other groups within the family led, and for the most part still lead, to quite large discrepancies. In the 60's of the previous century, the honeybadger
badger group, and this view is held, in part, to the present day. Those who do not assign it directly to the badgers subfamily Melinae in all events, closely affiliate Mellivora with Melinae. At the same time, the similarity between

was assigned

to the

!03

honeybadger and badgers is actually purely external, and expressed in general measurement, color type and, as is clearly assumed,
general appearance of the animal (see beyond).
It

is

likely that the viewpoint is

still

more widely accepted,

according to which the genus Mellivora forms a separate subfamily Mellivorinae close to the subfamily of badgers, Melinae, a

"neighbor" so to speak, to

it.

At the same time,

tails

as

was

in part

mentioned above (see also

beyond), according to skull, both in main features and

the subfamily of martens, true Mustelinae, and
is

many

de-

of dentition, the genus reveals greatest similarity to species of

strongly differ-

from the badger group, Melinae. The latter have different dentition both in structure and adaptive type, that is particularly expressed in structure of the posterior upper molar and in other characteristics (see beyond). According to the sum of all features, the genus Mellivora cannot and must not be united with the badger subfamily, Melinae, and it is very difficult to argue for its separation into an independent subfamily. Genus Mellivora must be considered as one of the typical, but specialized, forms of the marten subfamily, Mustelinae. This view of the position of this genus within the subfamilies was also previously expressed. Within the boundaries of the subfamily Mustelinae, the genus Mellivora is, to a certain extent, analogous to the genus of wolverentiated
ines,

Gulo.

If the

wolverine

is

actually a specialized deviant

gigantic marten, the honeybadger

may be considered

huge

1210
aberrant polecat, including also in this designation the above-men-

tioned

(179)

African genera of polecats^.

An

opinion was expressed about some genetic connection of

honeybadger with wolverine, but it is without serious foundation. Because of the closeness of the genus Mellivora to the remaining Mustelinae, the significant distance of the genus from Meles

and the subfamily Melinae

clearly considered to

in general points to the systematic position

of the skunk (Vonyuchka*) group, subfamily Mephitinae. They are

occupy an intermediate position between

it

"Mellivorinae" (honeybadger) and Melinae (badgers). However,

is

indubitable that, according to dentition for instance, skunks are

all

sharply differentiated from the honeybadger and

Mustelinae,

and are much more similar to badgers (Melinae). Therefore, the gap between g[enus] Mellivora and members of the subfamily of badgers, Melinae, is very sharp and deep. Nowadays, the subfamily of skunks is restricted in its distribution to the New World

Miocene-upper Pliocene) it was also Europe and in Asia. With all of this, the uniqueness of the genus Mellivora remains without doubt. Its independence is also shown by its relatively long period of existence from the middle Pliocene (Asia). The genus Eomellivora, nearest to Mellivora, is known from the upper Miocene and existed as several species during the course of the whole Pliocene both in the Old world (Eastern Europe Odessa district and Moldavia; Asia), and also in America. The early relationships and origins of both genera are unknown, but Eomellivora is evaluated not only as a quite typical form close to Mellivora but also as a more specialized genus than Mellivora in some respects. In general, the origin of the group is associated with the primitive Mustelinae (Thenius and Hofer, 1960). In the genus there is one species: Mellivora capensis Schreber,
only, but in the past (upper

found

in the

Old

in

1776.

The forms of
guishable.
In the

the genus are, apparently, practically indistin-

USSR,

distributed in the western part of

Middle Asia,

and associated with deserts and desert mountains. It has no practical significance (V.H.).
different forms, they are even

^According to their dentition which may consist of 28, 30 and 36 teeth in more strongly differentiated from the true Mustelinae than the honeybadger. *A Russian word literally meaning "stinking" animal or person Sci. Ed.

1211

804

RATEL OR HONEYBADGER3
Mellivora capensis Schreber, 1776
1776. Viverra capensis Schreber. Saugeth.
pp. 450, 588.
pi.

125. Also: 1777, 3,

1792. Ursus indicus.

Cape of Good Hope. Kerr. Animal Kingdom,

p.

188. India.

1851. Mellivora ratel. Horsfield. Cat.

India.

Mam.

E. Ind. Co., p. 120.

1920. Mellivora wilsoni. Cheesman. Journ.

27, p. 335.

Bombay

Nat. Hist. Soc,

Ram-Khormuz on

the Iran-Iraq boundary.

Diagnosis

Only species of the genus.

Description
In general proportions, appearance,
is

and behavior, the honeybadger

entirely unique and, in spite of the prevalent opinion, does not

It

resemble the badger.

polecat.

more quickly reminds one of a bear-cub on

it

the one hand, and on the other,

It

has

much

in

common

with the
its

often

moves

in

a sort of creep, extending

body,

back and raising its tail upright. Trunk quite heavy and massive, moderately elongated, tail short, limbs short but quite thick and strong. Head quite large and broad compared to body dimension, with blunt facial portion. Bare nose tip large, anteriorly flattened, black in color. Ear pinna greatly reduced and represented by short dermal ridges. Limbs with relatively large, broad feet, the lower surface of which is bare in winter and summer. Fore paws typically fossorial. Their claws very long, laterally compressed and moderately bent. Their distal halves are 805 light-horn, basal dark-horn. Length of longest claw 32-46 mm. Claws of hind limbs relatively large, but much shorter than those of fore limbs; their length 10-18 mm. They are darker, more massive and strongly bent, quite wide and almost completely covered by
its

bending

hairs of upper foot.

name, representing a translation of the among, Russian hunters in southern Turkmenia and it has the sense, "to use". This species is called by Turkomen "jf-a/i"), which means "dog-bear" (it dog). Description given mainly from material in ZMMU.
is

'"Medoed" [Honey-eater]

an

artificial

Latin generic name. Another

name

exists,

1212

804

Fig. 288. Ratel, or

honeybadger, Mellivora capensis Schreb. Sketch by A.N. Komarov.

805

Fig.

289. Ratel, or honeybadger, Mellivora capensis Schreb. Kushka

interfluve, southern

Kashan

Turkmenia, 1960. Photograph by Yu.K. Gorelov.


1213

Winter fur

is

long (hair length on posterior part of back 40-

50 mm),

closely-lying and consisting of coarse elastic, even some-

what bristle-like hairs. They are quite sparse (parting the hairs anywhere, one sees the skin), and without underfur, or it is very weakly developed. On sides, hairs noticeably sparser than on back; on belly, particularly on groin and axillae hairs extremely rare and here skin shows through strongly. Hairs on tail approximately of same length as on posterior part of back. Summer fur sparser and shorter hairs on back have length of

about 15

mm. On

belly, fur

still

sparser

it

is

half bare.

Skin very thick. Anal glands well developed. Teats, 2 pairs. belly, throat, chin, lower In winter fur, entire lower body

surface of

tail

and

its

terminal hairs, and also lateral surface of

body

side of head including ears, side of neck, outer surface of limbs as far as the middle of shoulder blade and middle of thigh is a shiny as well as anterior part of muzzle including eye region

black color. Sometimes, the fur has a very light brownish tinge. On top of head, a little behind the eyes, a white area begins,

having an arched anterior border and covering the whole upper body. This runs over the upper neck as a narrow band, in width corresponding with the area on the head, sometimes even a little narrower, gradually broadening in the shoulder region and reaching
its

greatest width in the middle of the trunk or nearer the

thighs.

At the tail base, this area strongly narrows and extends as a narrow band occupying the whole upper side. The hairs on the very end of the tail are black. On the whole, it has a typical dorsal saddle pattern extending, however, to the neck, head and the upper
side of the
tail.

In a stretched skin, the light area has a pear-shaped

tail.

or flask-shaped form with a narrow "extension" on the

the dorsal saddle

806

The general tone of this area is is somewhat variable. Hairs on

white, but in details of color,

the head and

upper neck are usually white to their base. Sometimes, however, their bases are somewhat darkened and they are slightly visible against the general white background. Hairs white to the base, or with weakly developed darkening at the roots form a narrow band edging the dorsal saddle from shoulder to thighs. The remaining

main

part of the saddle patch

is

covered by hairs with a white

less

distal half

and with a wide dark base. These bases more or

(usually strongly) shine through, and the whole area has a unique

color a mixture of pure white and dark tones resembling the color

1214
of gray with remains of black hairs ("salt and pepper"). The white
tone
is

cold and only sometimes has a light ocherous film.

The

above-mentioned edge band of pure white hairs is sometimes, especially with a strong admixture of dark in the saddle patch, is separated in the form of a fuzzy lightening on the general background, and sometimes the edge of the saddle patch also has the

same color as in the middle part. The summer fur has the same color as in winter, but the color of the light dorsal field is more intensively mottled with dark, with
the pure white areas on the neck and head absent

they are of the

in its poste-

same color
als,

as the

middle of saddle patch. Sometimes

rior part, a lighter

narrow border

is

noticed. In different individu-

the dark tone

less,

is

visible through the white,

is

sometimes more and

sometimes

and the general color

lighter or darker but these

differences are, apparently, small.

Sexual differences in color are absent, and age differences are

undescribed.
(Africa).

Among

the exotypic variations,

malanism
is

is

known

The geographic

variation in color, if present,

very weak.

Skull relatively large and quite heavy, of entirely unique structure'*.

The upper

profile of the skull presents itself as a quite even,

its

arched line having

highest point in the parietal; quite quickly

descending posteriorly, and very gently sloping forwards without

any rise in the interorbital region. The profile of the nasal opening, which is greatly displaced backward, serves as a continuation of
this line

with a small projection. The facial part of the skull,

is

al-

though wide,

short; its length is

approximately two times shorter

long and very narrow

than the braincase.

The
is

postorbital constriction

is

its

width

always less than that of the interorbital area, usually considerelongated and wide, ably so (by 5-10 mm). The braincase is large

broadening posteriorly and swollen

occipital crest is strongly developed,

in

the parietal region.

The

weak (much weaker than

in

and the the badger) even

sagittal is relatively
in the

very old aniof

mals. Zygomatic arches are strong but not very massive, in the

medial portion arching upwards very strongly. The

processes have wide bases, but are short.

lateral lines

the interorbital region are parallel or almost parallel, the supraorbital

is often compared to the skull of the badger as the was based on misunderstanding. One may see similarity only in general dimensions and known massiveness; generally the skull of the honeybadger is more quickly recognized as the skull of an enormous marbled polecat.

"The skull of the honeybadger

similar. This

most

1215

807

Fig. 290. Skull of

honeybadger, Mellivora capensis Schreb.

The mastoid processes

are developed very strongly, situated

almost horizontally, and strongly projecting to the sides, whence

1216
the mastoid width of the skull
least equal to
it.

is

greater than the zygomatic, or at

Mastoid processes are flattened, and dorsally even somewhat concave. Their lower surface fuses with the upper wall of the auditory tube^ and the posterior part passes to the occipital
crest.

The

paroccipital (lateral occipital) processes are large, sepa-

rated both

from the occipito-articulars and also from the auditory

bullae and greatly projecting ventrally and

somewhat

externally.

Bony
808

auditory bullae very large, swollen, angular-oval and

strongly diverging anteriorly

their anterior walls adhere closely and partially fuse with inner parts (those directed towards sagittal

plane of skull) of postarticular processes. Hard palate short and

wide;

its

interior width

that) than distance

between molars 2.5 times less (or about from posterior median sphenopalatine notch to

alveoli of middle incisors. Portion of palate lying behind toothrow

and broad. Infraorbital foramen small, its diameter not more than half diameter of upper canine alveolus. Dentition strong and massive with short apices, wide at base.
relatively short

Carnassial teeth particularly massive, having large main cusp with

wide base and well-developed, quite wide secondary inner blade. Upper molar quite typical of martens and polecats small, strongly

elongated in transverse direction and

its

transverse diameter al-

most double longitudinal (anteroposterior). Inner blade considerably greater than outer; they are separated by a marked isthmus. Canines strong and massive, broad at the base, but quite short. Anterior upper premolar relatively large, having two roots. In con'

trast to that

observed in previously described species of subfamily

is

Mustelinae, and also in badgers and, in part, in otters, there

no

tendency towards reduction or loss of

Sexual differences
able,

this tooth.

in skull structure are

absent or not notice-

and no age variation has been described.

has, in terminal quarter, a

The OS penis
evenly divided

bend upwards

tip

located nearly at right angle to axis of proximal half. Tip itself not

one "horn"
is

is

larger than the other (Pocock, 1941).

In general, structure

typical for martens

and polecats (subfamily

Mustelinae) and differs strongly from that characteristic of badgers.

The body length of males

680 mm;
tail

(2) is

length of males

is is

length of hind leg in males

'It

680-750 mm, of females (1), 180-205 mm, of females, 172 mm; 120-125 mm.

is

indicated (Pocock,

1941) that "the cavity of the long swollen auditory

capsule posteriorly opens into the hollow of the mastoid process".


1217

Condylobasal length of male skull (3) is 139.5-145.7 mm, of (1), 130 mm; zygomatic width of males (3) is 80.0-86.2 mm, of females (2), 75.2-80.3 mm; mastoid width of males (4) is
females
82.8-86.4
(4) is

mm,

of females, 80.7

mm;
(2),

interorbital width of

males

31.7-33.8

mm,

of females

32.0-32.6

mm;

postorbital

width of males (4) is 23.6-27.5 mm, of females (2), 27.7-30.5 mm; width of muzzle above canines in males (4) is 32.5-34.6 mm,
of females, 30.8
in

mm;
is

length of upper toothrow (without incisors)

males

(4) is

38.5-40.2

mm,

of females (2), 30.8-37.2

mm.

Weight of male
female, 9.1 kg.^

12 kg (September) and 16 kg (October), of

There

is

sexual dimorphism in body and skull measurements

the females are smaller than the males

but
is

they are relatively

insignificant. Difference in overall weight

more obvious (V.H.).

Systematic Position

Only species

in the genus.

Geographic Distribution
Africa, a large part of

range of the ratel

in the family.

is

Near Asia, India, western Middle Asia. The one of the most extensive of the species ranges

809

Geographic Range

in the Soviet

Union

The range

is

not great, and constitutes a very small part of the

species range

its

extreme northern

part,

and occupies an insig-

nificant part of the country.

The southern border of

the range within the boundaries of

USSR

coincides with the state frontier from the Caspian Sea to Amu-Dar'ya. Occurrence and places of permanent occupation of
*From material of Z[oological] M[useum of] M[oscow] U[niversity] from Turkmenia, collected by A.M. Shukurov, I.V. Zhernovov, V. Svyatoi, S.I. Bil'kevich, E.I. Shcherbina, A.N. Sukhinin and from one specimen collected by Varentsov (Ognev,
1931).

Measurements of Indian animals (Pocock, 1943) are: body length of males (5) is 690-725 mm, of females (2), 625-650 mm. Condylobasal length of male skull (6) is 131-141 mm, of females (6), 121-132 mm; zygomatic width of males (8) is 74-80 mm, of females, (5), 71-76 mm. Weight of male (2) (time [of year] is not mentioned) 9.7-10.4 kg, of females, 7.7 kg. Data relate to both Indian subspecies. The maximum body length of Indian animals of 82.5

cm

(Novikov, 1956)

is

apparently a misprint.

1218

809 Fig. 291. Limits of distribution of honeybadger, Mellivora capensis Schreb. in the USSR. Dots indicate several separate habitats in Karakum and other deserts of Turkmenia. Many known occurrences of the animal along boundary of range between Amu-Dar'ya and Caspian Sea not plotted. Data of 1962. V.G. Heptner.

the rate] are

known along

the Atrek [river], along the entire Kopet-

Dag

Kopet-Dag plains, along the Tedzhen and the Gyaz'-Gyadyk mountains (right bank of upper Tedzhen), throughout the whole Badkhyz (expanse between the Tedzhen and Kushka) and in the highland area between the Kushka and the upper Murgab. Apparently, the honeybadger also exists in the area from the Murgab to Amu-Dar'ya it was found on the left bank of the upper course of this river in Khalach region.
[range] and nearby

1219

The western border on

coast, to a little north of

the north

is

formed by the Caspian

Krasnovodsk. Thence or slightly further north (south shore of Kara-Bogaz-Gol), the northern border of the range begins. It passes to Kazakhla village in the southern Ustyurt Chink (in the place where the borders of Turkmenia, Kazakhstan

and Karakalpakia meet) and Kaplankyr and farther along the Chink no (Aganash, Kurgankyr, Zangibaba), occupies Sarykamyshsk depression, and through Deukal village (Deukaskenkala, Onkhauz and
Doudyr), generally
at the latitude
it

of Kunya-Urgench, and some-

what
itself

to the

west of

reaches the

Amu-Dar'ya

delta. In the delta

and

in its

contiguous cultivated lands,

in particular the

Khiva
the

honeybadger is Kunya-Urgench district, the border turns

oasis, the

not, apparently, encountered.

From

to the southeast,

becom-

ing the eastern border of the range and, apparently passing around

Khiva

oasis,

state border.

recorded

at

and going farther along the Amu-Dar'ya, reaches the In the middle course of the river, the ratel was Darganat (tugai) and in the upper course, it was
at

recorded, as noted above,

Kalach near Kerka. The honeybadger has not been recorded on the right bank of
is,

Amu-Dar'ya and
from the south

apparently, absent there.

The Amu-Dar'ya

constitutes an insurmountable barrier for this species dispersing

from Iran

and Afghanistan.

Inside the outlined region, the ratel has been noted in a series

of places, not only in clay deserts along the chinks of the desert
plateau, in declivities in mountains, river valleys "pistachio

savanna"

etc.

but also in sandy deserts, both with saxaul and with-

out. Apparently, the distribution of this species within the borders

of our country

is

more or

less continuous, although its

numbers

evidently predominate to the south, and in areas with a more or

less interrupted relief

and with more compact

ratel is

soils.

Within the
for

boundaries of the described region, the

known,

exam-

ple, from the following places: Sharlouk in the south of western Turkmenia, a little north of the Atrek river and west of Karakal; sand 60 km north of the railway station Geok-Tepe; Bala-Ishem well in the Usboi 150 km north of Kyzyl-Arvat railway station; Bakhardok well between Ashkhabad and Sernyi zavod [factory]; sands of Tedzhen region; lolbarsli well (north of 38 N. lat. and a little east of the 60 E long.); railway station Uch-Adzhi between

Mary [Merv] on

the

Murgab and Amu-Dar'ya

(sands);

Darvaza

1220
(northwest of Sernyi zavod) Kyrkkaya (100

km

east of Darvaz),

Zaunguzsk Karakum, and

tury,

others'.

Beyond that, in the beginning of the 90s of the previous cenwhen the first information on the occurrence of honeybadgers
at all
it

within the borders of our country appeared (Bikhner and Zarudnyi,

1892; Varentsov, 1894) there was no information

a long time, and the prevalent opinion
sionally

about

it

for

was

that

appeared occa-

any event, represented an extreme rarity. New data appeared only in 1918 (Bil'kevich, 1918) and in the 20's (Ognev and Heptner, 1929). From these, it was clear that the honeybadger, although rare, lived permanently in the Kopet-Dag, and across the directly adjacent plains in extreme southwestern Turkmenia from the Tedzhen to the Atrek. It was later elucidated
us, or in

among

that

it

occurred
in

at

furriers in these places

starting

from 1924,

although

small numbers (individual animals). Suppositions con-

cerning the great rarity and extremely limited range of the ratel
with us persisted until recent times (Novikov, 1956
"exclusively southern Turkmenia").
Investigations in the 50' s and the very beginning of the 60'

"very

rare",

and, to a lesser extent, in the 40' s,

all

showed

that the ratel is not at

rare in the country, being distributed northward to the Ust'yurt

and occupies the whole Karakum. It was shown to be not rare even at the extreme northern border of its range in the USSR, and is quite common in the south. Apparently, one may think that at present the honeybadger occurs more often in Turkmenia than the badger [Meles] and is more widely distributed. It is significant that
811

in Turkmenia, at least in the southwestern half of the country, the honeybadger has a special popular name. Russian hunters also have

a special designation for

it

(see above).

Undoubtedly, the present picture of the range and population

of the described species, appearing completely unexpectedly, the

wide zoological investigations which have unfolded Turkmenia in the last ten years. However, there was also an increase in numbers of the species at least in the south, where it undoubtedly became much more abundant than in the first quarter
result of
in
''The honeybadger is found in a great number of places along the abovementioned southern frontier of our part of the range. They are not listed. Range according to Bil'kevich, 1918; Ognev and Heptner, 1929; Sukhinin and Shcherbina, 1955; and unpublished material of V.G. Heptner, and mainly Yu.F. Sapozhenkov, Yu.K. Gorelov, I.V. Zhemovov, I.V. Svyatyi, V.P. Kostin, and collec-

tions of Z[oological]

M[useum

of]

M[oscow]

U[niversity].

1221
of our century, and the borders of its range shifted to the north. Nothing be said about the extent of its increase in numbers and the period during which it took place. In any event, the present north-

ernmost habitats
E. long.) to

lie at a

650

km
to

distance ranging from 350 km (at the 56 (along longitude 59 E. long.) from the pre-

(lO's and 20' s) and the more eastern parts 800-850 km. Settlement proceeded both from the direction of Afghanistan and from the direction of Iran, but was probably more intensive in western Turkmenia and thence into

viously

known ones

even farther

up

its

northern and northeastern parts. Despite being a large eurytopic

it,

species, sandy deserts are for

seemingly

less favorable.

Geographic Range outside the Soviet Union

In Africa, the

honeybadger

is

distributed at present

from the ex-

treme south (Cape region) northwards to Ethiopia, northern Somali and Sudan (Suakin near Port Sudan on the Red Sea). In the past,
it

is

evident that

it

also occupied the regions adjacent to the Nile

Egypt and reached the Mediterranean Sea in this part of Africa. However, the northern border of the range in Africa is quite indefinite. Judging by features of the species described from the Trans-Caspian part of the range, in the east, it apparently passes
in

generally keeping to the 20 N.

lat.

along the southern edge and

it is,

in

the southern parts of the Sahara, and in the west,

more northerly

along

apparently,

the hills of the

its

Ennedi desert, Tibesti, Adrarlevel).

Iforos and others. In any event,

presence was established on the

Air plateau (Asben, 1800

the Sahara to southern

above see
(Ifni,

Thence, the border

the western parts of

of the range passes in an unclear

way through

Morocco
its

Sus and the Valley of

Um-

er-Ryabii, a

little

south of Casablanca). In the Sahara, the ratel

northeastern parts, notably the Libyan

only avoided probably,

desert.

The range probably does not include the inner parts of the forested region of West Africa, although it has been recorded in the Congo (Ituri region).
In Asia, the range includes Syria, Palestine and the Arabian

Peninsula, Iraq (except, probably, the northern parts), the greater

part of Iran^ Afghanistan (except, probably, the higher parts of the
'Missone (1959) refers to the range of ratel it may, however, exist author, honeybadger inhabits only a small territory
incomplete, believing that
in Iran as

very limited and clearly

of the country (in

According to this extreme southwestern corner Khuzistan) between the head of the Persian Gulf and the southern contd. on next page
in other places. in the

1222

Hindu Kush),
to

inces to Bengal inclusive (absent in

from the former Northwest Frontier ProvAssam and Burma), northward the foot of the Himalayas and the southern zone of Nepal, in the
India,

south to the sea coast

It is,

to the southern extremity of the peninsula.

it

apparently, absent in the Malabar coast. In Ceylon,

it

is

ab-

sent. In the described reconstructed range

has apparently not

undergone essential changes, except

in

Egypt (V.H.).

Geographic Variation
With so
significant a range, the ratel undoubtedly displays
it

known
Thus,

geographic variation; however,

the differences

is

not, apparently, great.

between the two "polar" forms

the South African

capensis and the Indian indica

lie

in

insignificant differences

in color and in the somewhat smaller average dimensions of the Indian subspecies. The two Indian subspecies differ insignificantly

from each other only

in density of the fur

(Pocock, 1941).

About 15 subspecies in all have been described, almost all from a single specimen. There is no special revision of geographic
variation in the species.
812

Within the boundaries of the USSR, there

apparently, not differing in any substantive

is

only one form,

the Indian.

way from

Indian honeybadger, M.

indica Kers, 1792.

For characteristics of this form, see above. Western part of Middle Asia northward to the southern Chink
of Ustyurt, and eastwards to the Amu-Dar'ya.

Outside the

USSR in

Afghanistan, Iran (except, perhaps, the

southwestern), [West] Pakistan and the western part of India (not

east of southern Nepal).

Two

forms are usually considered

indica and inaurita.

to exist in the territory of

Hindustan
hairs are

They

in that in the

second (eastern) the fur

It is

found on the heel.

in the

from each other only somewhat denser, and assumed that this form may also
differ
is

be found
contd.

provinces contiguous to Afghanistan (Pocock, 1941)

from previous page

half of the border with Iraq.

belt along our borders

On

the other hand,

to the

from Tedzhen

its range was described Gorgan (Astrabad) Gulf on

as a

narrow

the Caspian

Sea. Judging by the distribution of the species in the

countries and by
its

USSR

and
it

in

neighboring

biological characteristics, one

may

allow that

exists over the

whole of Iran, except probably the northwestern, maybe, the wooded region of the Elbrus mountains (it was, however, found near Gorgan city; Bil'kevich, 1918) and the vast deserts of the middle and eastern parts of the country (Dash't-e-Kavir, Dash't-eLut). Proceeding from these data, the range is given in the map (Fig. 287).

1223

although this

is

not quite understandable (Pocock's

text

is

contradictory). Since our honeybadgers live under

ditions (winter) than those under

which the
is

more severe conIndian form lives, it is

possible that the fur of the former

i.e.
it

denser than that of true indica,

is

like inaurita.

tween these forms are

mals.
is

However, the differences themselves bedoubtful: our museums have no materials of

these races and therefore the

name given

is

retained for our ani-

The

relationship to the form wilsoni from southwestern Iran

this

perhaps unclear, but judging from everything,

form, de-

scribed from individual characteristics of one specimen, does not

have substantial

truth.

Nepal and of range of M.c. southwestern M. pumilio Pocock, 1946 Hadramawt, southern Arabia; M. leuconota 1867 West Africa northwards southern Morocco; M. signata Pocock, 1909 Leone; M. consica Thomas Wroughton 1907 Lake Chad; M. buchanani Thomas, 1925 1910 Air (Asben), Sahara; M. abyssinica M.c. broekmani Wroughton, 1920 Somalia; M. Lydekker, 1906 Congo; M. makwelli 1910TanganThomas, 1923 Kenya; M. sagulata M. vemaj/ 1932 Bechuanaland; M. capensis Schreber, 1776 South and Southwest
Hodgson, 1836
parts

The following forms are usually described from parts of the USSR. Independence of a series of these forms, including Palearctic ones, is doubtful. 1) M. inaurita
range lying outside the
the
east
it^
;

2)

wilsoni Cheesman, 1920

Iran, Iraq (?); 3)

c.

4)

Sclater,

to

5)

Sierra

6)

et

7)

plateau

8)

c.

Hollister,

Ethiopia; 9)
cottoni

10)

Ituri

region,

11)

12)

Hollister,

yika; 13)

Roberts,

14)

Africa.

The

actual
is

number of forms of honeybadgers

clear at least

is,

evidently,

smaller. This

from comparison of the places whence

several forms were described (abyssinica-broekmani; leuconota-

signata; consica-buchanani; indica-wilsoni) (V.H.).

Biology
Population. Everywhere within the boundaries of the range in the

USSR,

it

is

not often encountered, but

it

is

not particularly rare.

Numerical indices of population do not

exist.

'The suggestion of "Northwestern Frontier Provinces", i.e. contiguous to Afghanistan (Ellerman and Morrison-Scott, 1951) is hardly correct. It contradicts the main data of Pocock's monograph (1941) and other indications of these same authors
(distribution of the

form

indica).

1224
Habitat. Within the limits of natural landscapes in
the
in
its

range,

honeybadger may

be called a eurycious species.

It is

met with

sands of the Karakum; in sands and foothill plains of the Kopetrange; in montane canyons of the latter;

Dag
hills

among

steppes and

of Badkhyz; in marsh

tit*

plains covered with saxual, Russian

thistle [Salsola]
813

and sagebrush; among deep ravines of the south-

ern precipice of Ust'yurt [plateau] with stunted vegetation of shrubs,

sagebrush, and succulent saltwort; in cultivated river valleys

the bai** landscape; and in dense riparian tugai thickets.

among

Food. Insufficiently studied. In India,

bees,

it

feeds on small ro-

dents, birds, reptiles including cobra, as well as grasshoppers, miner

wasps and

their larvae
Its

attack poultry-yards.

bias towards

and honey. In India, they sometimes honey is noted in South Africa.

iX^.

Fig.

292. Habitat of honeybadger at northernmost range border. Precipices

bio-

tope of arkhar [O.

ammon]. Foot of southern Chink,

Photograph by V.P. Kostin.

Ust'yurt, Kazakhly village.

*The Russian word {pukhlyakoba) means marsh

a descriptor of landscape
is

tit,

but

its

significance as

unclear
Ed.

Sci.

Ed.

**bai
i.e.,

is

a Turkic

word
Sci.

referring to a wealthy

landholder

in

Middle Asia;

river-valley land

1225

According

to data collected in the

most recent period

in the

USSR, the honeybadger may be called an omnivorous animal. It eats plant food in captivity; in nature, this event is unknown. In its food,
remains of great gerbils occur repeatedly, and
a case of eating the carcass of a
it

also eats hedgehogs;

lamb was observed (Kostin,

tiles,

Sapozhenkov, Gorelov, Zhernovoi and Svyatoi, 1963). Among repfeeding on the desert monitor was recorded in Badkhyz (Sukhinin
and Shcherbina, 1955 and others). It very often feeds on Horsfield's which it digs out even in winter. Small lizards and round-head
its

tortise

lizards [Phrynoce-phalus] also occur in

food.

At the southern edge

of

Ust'yurt

(V.P.

Kostin),

up

to

Phrynocephalus were found

in feces.

30% reticulate scales of Among insects, grasshoppers

were found,

in particular the saxual

humpbacked grasshoppers, and

darkling beetle (Sapozhenkov, Kostin, Zhernovoi and Svyatoi, 1963).

Home
km; by
it

range. Information

is

lacking.

On

the edge of the Ust'yurt,

less than 2.5

tracks of honeybadger were traced for a distance not

follows that, the

home

range

is

quite large (V.P. Kostin).

lives in

Burrows and
itself.
It

shelters.

The honeybadger
is

burrows dug

digs very well, and

able to dig a tunnel into very

hard ground in 10 minutes. The honeybadger usually digs simple burrows with one passage and nesting chamber. The dimensions of the latter: 37 x 43 and up to 60 cm. Bedding does not occur in the nesting chamber. The length of the burrow is not large: from 1 to 3 m. The diameter ranges from 17-18 to 30 cm, more often 25-30 cm. The distribution of burrow depth ranges from 25 cm
?14

to 1.5

m. Apparently also occurs

in

rocky shelters. In Ust'yurt, an

accumulation of feces

a "latrine"

was

found

in a small

cave

among limestone blocks

(V.P. Kostin).

Daily activity and behavior. The honeybadger has no regular

515

rhythm of daily activity. As a rule, it leads a crepuscular-nocturnal way of life (Sapozhenkov et al.; 1963; V.P. Kostin), but is sometimes active during the day, especially in March-April, on warm, sunny days. Its daily activity has been observed in India. In South
Africa,
it

is

usually active during the day.

Seasonal migrations and transgressions. Information is absent. Reproduction. Until recent times, nothing was known about
reproduction, except that in South Africa, a female gave birth to

two cubs

after a

6-month pregnancy.

Signs of rut were noticed in female in Ashkhabad

Zoo

at the

end of September.

male caught on 10 October, had large testes. In Ashkhabad Zoo, a female caught on 6 May delivered on

1226

X
^A
^

814

honeybadger in the northern limit of its distribution in northern Karakum. Arkhar also live here and rarely cheetah and leopard. Kaplan Kyr village, southern Chink, Ust'yurt. Autumn 1962. Photograph by Yu.F. Sapozhenkov.
Fig. 293. Habitat of

!14

Fig.

294. Hilly landscape in

ratel

or honeybadger.

Badkhyz preserve, southern Turkmenia. Habitat of May, 1961. Photograph by Yu.K. Gorelov.

1227

\-0

-fO

-10

CM

k^

Fig. 295. Tracks of honeybadger. Sketch

by V.P. Kostin.

1228
birth to one cub. In the first days of May, a and cubs with erupted teeth and opened eyes was observed. The duration of pregnancy, therefore, is about 7 months, which is close to the South African data. Growth, development, and molt. Information is lacking. Enemies, diseases, parasites, mortality, competitors, and population dynamics. Data are absent. Field characteristics. The color of the honeybadger masks it very well in solonchaks. Moving in an uneven gallop, it is very

31

May; she gave

lactating female

816

reminiscent of the general form of the wolverine. Prints of

sausage-like form, 4-7
angry,
it

its

tracks are wider and shorter than the badger. Its feces have a

raises

its tail

In addition to

cm long and 12 mm diameter. Becoming upward when walking (V.P. Kostin). the gallop, by which the honeybadger can proit

ceed for a prolonged time,

a hoarse "khrya-ya-ya-ya".

also walks and even half-crawls,

sprawling on the ground. The voice of the honeybadger resembles

At the time of

rut,

the male

loud grunting sound (Sapozhenkov

et al., 1963).

makes a The honeybadger

climbs trees well.

It

gives off a frightening smell and exceptional

movement of

the skin surface (P.Yu.).

Practical Significance

Due
places,
cially
it

to

its

small numbers and because

it

inhabits little-populated
It is

has almost no practical significance.

is

not hunted espe-

and

for the

most

part captured accidentally.

this,

Honeybadger

are

sometimes caught alive for zoos. For

1963). (P.Yu.).

the animal's tracks are

al.,

followed and they are dug out from burrows (Sapozhenkov et

Subfamily of Badgers
Subfamilia

MELINAE

Burmeister, 1850

Genus of Badgers Genus Meles Brisson, 1762

1762. Meles. Brisson. Regn. Anim., 1758.
p.

13.

Ursus meles Linnaeus,

1229
1795. Taxus. Cuvier et Geoffrey. Mag. Encyclop.,
2, p. 187.

Ursus

meles Linnaeus, 1758. 1925. Meledes. Kastschenko. Zap. Fiz.-mat. vidd. Ukr. akad. nauk, I, No. 4. Meles leptorhynchus Milne-Edw. Described as a
subgenus. (V.H.).

Dimensions

large.

General conformation heavy and clumsy; body relatively short,

considerably wider posteriorly than anteriorly; head relatively small;

Limbs not shortened, massively plantigrade with long claws of digging type; webbing between digits absent. Ear pinnae
tail

short.

normally developed. Hair cover coarse, long but sparse, with weakly

developed underfur. Coloration from combination of black and white, type of saddle-patch developed to an extreme degree. Between the anal opening and. root of tail there is a glandular "pocket", into which an odorous secretion exudes. Anal glands well-devel2 abdominal, 1 inguinal. oped. Teats 3 pairs Skull quite massive and heavy, but not broad, moderately high.

Braincase not of increased volume, facial part of skull elongated

and narrow

(its

length

more than

its

width), that part of hard palate

lying behind toothrow long and relatively narrow, mastoid proc-

esses well developed but not very large, paroccipital processes

well-defined, but short. Mastoid width of the skull

less

is

considerably

than zygomatic. Auditory bullae of moderate size and not

swollen. Auditory tube not covered by mastoid process and opens

independently

in front
is

of

it.

Dentition

a specific type, fundamentally differing

It

from

that

characteristic of Mustelinae.

represents a combination of teeth of

a sectorial type with those having clearly defined structural features of a tuberculo-grinding (crushing) tooth.

The

structure of the

dentition

is

one of the most characteristic features of the sub-

family.

It is

(Lutrinae).

more Upper

similar to the dentition of the subfamily of otter carnassial tooth not largest in toothrow

it

is

short and triangular in form.

larger than carnassial,
flat,

Upper molar

is largest,

considerably

tuberculate and of grinding type.

Lower

carnassial tooth elongated. All cheek teeth,

are of

except upper molar,

more

or less distinctly sectorial type.

Complete dental formula

individual first premolars or

is

I-C-P M = 38.

14

However,

all

of them are frequently lost and


1230
the formula changes correspondingly (see below, description of

species).

omnivorous forms, feeding mainly on invertebrates and small vertebrates. Ecologically they are entirely flexible and are encountered from closed taiga and broad-leafed forests to steppes and semideserts, and in part true deserts; they do not avoid mountains and are met with at great heights. Digging capability is strongly developed. There is a latent stage in development of fertilized eggs, and winter sleep. The range of the genus is vast and occupies Europe, a considerable part of Siberia, Near, Central and Eastern Asia as well as

They

are

Middle Asia.
In the current century, several species

were included

in the

genus other than the European M. meles Cretan M. arcalus, Asiatic M. leptorhynchus, Tibetan M. leucurus and Japanese M.

anakuma. For our fauna, two species M. meles and M. leptorhynchus were accepted. In actuality, separate species must not be distinguished in the genus. Even the extreme races, as regards their distribution and their characters, as well as their form are interrelated. The differences between them, as between M. m. meles and M. m. amurensis, may be very great in color and even dentition (in the European badger, there are as a rule 38 teeth and in

amurensis

34).

The

circle of genera in the subfamily includes besides Meles,

(American badger), Arctonyx (Indian "sand" badger), Helictis (including Melogale:* "ferret" badgers of southMalay archipelago). All of them eastern Asia) and Mydaus (teledu are sharply distinguished from each other (Helictis is sometimes
the genera Taxidea

separated in an independent subfamily) and

lish their systematic relationship.

it

is

difficult to estab-

The genus Meles may

be, appar-

ently, considered in this sense a little specialized form, although

its

dentition

is

in

some

respects (development and relationship

between upper molar and carnassial) more specialized than in some other forms. Apparently, Meles stands closest to the American Taxidea, to which it is most similar in general appearance. Taxidea must, however, be considered on the whole and particularly in its
skull, a

more specialized form, although

it is

Meles as regards specialization of the

dentition.
is

somewhat inferior to The southern Asiatic

Arctonyx, also closely related to Meles,

*This
is

usually considered a

now

the accepted

name

for the

genus of ferret badgers

Sci. Ed.

1231

518

Fig.

296.

Range of

the genus of badgers, Meles Briss., and species range of the

common

badger, Meles meles L. Borders of the range in Afghanistan, China and, in

part, in the

Indochinese Peninsula are to a certain extent given approximately. Dotted

main race groups of badgers European meles, Siberian Chinese "sandy" badgers, arenarius-leptorhynchus and Far East badger, amurensis-anakama. A question mark in the region of the watershed of Kama and Pechora points to the unclear limit between the two race groups. V.G. Heptner.
lines separate the regions of the distribution of the

more specialized form (Pocock, 1941), although, apparently, without sufficient basis.

Very primitive representatives of the subfamily, or forms close {Broiliana, Stromeriella) are known from the early Miocene of Europe and Asia. A series of genera which indicate the great
to
it

1232
variety of forms of the group are

known from

the upper Pliocene

of the Old World. The majority of them, however, as regards the

line leading to the

genus Meles, represent

lateral

branches with

various specialization {Palaeomeles, Plesiomeles, Taxodon and

others).

Aside from them, Parameles (Odessa), Melodon and

818

Parataxidea stand closest of all to the genera Meles and Taxidea. Parataxidea is known from the upper Miocene and lower Pliocene
of Asia and
is

widely distributed,

itself

representing a separate,
the source

more

specialized, group.

Melodon must be considered

of the genus Meles.

The genus Meles itself appeared in the upper Pliocene from whence are known M. gennevauxi, M. taxipater, and M. thordi and from which through M. m. atavus the line leads to modern M.
meles. This
is,

apparently, the typical Asiatic form, developing

without close connection to the American badger (Taxidea) which

is

known only from America (upper

The genus includes one
species:

Pliocene).

It

is,

apparently,

related to the Asiatic Parataxidea.

Meles meles Linnaeus, 1758.

little

In practice, the forms of badger are of

positive impor-

tance (fur, destruction of

819

some

pests).

In the

USSR

it is

distributed throughout the greater part of the

European territory of the country, through the southern half of Siberia and the Far East, and in Middle Asia (V.H.).

BADGER
Meles meles Linnaeus, 1758
1758. Ursus meles. Linnaeus, Syst. Nat. Ed. X,
1, p.

48. Uppsala,

Sweden. 1785. Meles taxus. Boddaert. Elench. Animal, 1816. Meles europaeus. Demarest. Nouv. Diet.
tute for meles.

1,

p. 80.

Europe.

Hist. Nat. Substi-

1844. Meles anakuma. Temminck. Fauna Japonica,

pi.

Mamm.

p.

30,

6.

Vic. Nagasaki, Japan.

J.

1847. Taxidea leucurus. Hodgson.

763,
pi. 29.

Asiat. Soc. Bengal,

16, p.

Lhasa, Tibet.

1859. Meles taxus amurensis. Schrenck. Reisen und Forsch.

im

Amur-Lande,
[rivers].

p. 17, pi. 1, Fig. 1.

Amur

near mouth of Ussuri

1233
1867. Meles leptorhynchus. Milne-Edwards. Ann. Sc. Nat. Zool.,
8, p.

374. Vic. Peking [Beijing], China.

1875. Meles canescens. Blanford. Ann.

Mag. Nat.

Hist.,

16,

p.

310. Abadekh, between Shiraz and Isfahan, Iran.

1891. Meles schrenkii. Nehring. Sitzungsber. Ges. Naturf. Freunde

Berlin, p. 103. Substitute for amurensis Schrenck.
1, p. 111. Rynbetween lower Volga and Ural [rivers]. 1899. Meles meles typicus. Barrett-Hamilton. Ann. Mag. Nat. Hist., 4, p. 384. Substitute for meles Linnaeus. 1900. Meles taxus sibiricus. Kastschenko. Kashchenko. Opredelitel' Mlekop. Zhiv. Tomskogo kraya, plate 15. Former Tomsk

1895. Meles taxus arenarius. Satunin. Arch. Naturg.,

peski,

governance. Restricted to (Stroganov, 1962) Kolyvan', about

400 km north of Novosibirsk. 1901. Meles amurensis altaicus. Kastschenko. Kashchenko. Ezhegodn. Zool. muzeya I. Ak. nauk, 6, p. 613. Southern end of Teletsk Lake, Altai. 1901. Meles amurensis raddei. Kastschenko. Kashchenko. Ibidem.,
p.

613. Steppes of Trans-Baikaliya.

i

1905. Meles meles minor. Satunin. "Priroda

okhota", No. 2. Borzhom, Trans-Caucasus. 1907. Meles blanfordi. Matschie. Wiss. Ergebn. Filchner Exp. nach

China, 10,

1,

p.

143. Kashgariya.
.

1910. Meles tianschanensis Hoyningen-Huene. Zur Biol.

d. estland.

Dachases,

p. 63.

Tyan'-Shan' [Tien Shan].

J.

1913. Meles melanogenys.

32. p. 433.

Allen. Bull.

Amer. Mus. Nat.

river,

Hist.,

Musan on Tumangan

northern part of

Korean Peninsula.
1916. Meles meles ponticus. Blackler. Ann. Mag. Nat. Hist., 18,
75. Skalita near
p.

Trabzon (Trapezund), [Trebizon], Asia Minor.

inst.
p. 50.

1926. Meles meles caucasicus. Ognev. Uchen. zap. Sev-Kasvk.

kraevedeniya,
forestry

1,

Vic. Vladikavkaz.
1,

1926. Meles meles tauricus. Ognev. Ibidem.

p.

51. Beshuisk

camp

in

Chatyr-Dag, Crimea (Crimean preserve).

478. Southern slope of Talassk Alatau,
775. Aleksandro-

1931. Meles leptorhynchus talassicus. Ognev. Zveri Vost. Evropy

i

Sev. Azii, 2,

p.

western Tyan'-Shan' [Tien Shan].

1931. Meles meles heptneri. Ognev. Ibidem.,
p.

Nevskaya north-west of Kizlyar, eastern Cis-Caucasus. 1940. Meles meles severzovi. Heptner. Zeitschr. f. Saugetierk.,
p.

15,

224. Arkit, south of Syry-Chilek Lake, south of the north-


1234
eastern extremity of the Chatkal range, Tyan'-Shan' [Tien

Shan].
1962. Meles meles aberrans. Stroganov. Zveri Sibiri,
2,
p.

167.

Akmolinskaya

district,

Bogembai

village,

northern

Kazakhstan (V.H.).
820

Diagnosis

The only species of

the genus.

Description
General body constitution heavy and massive, especially
the broad
in full

winter pelage. Entire body seems to possess a wedge-shaped form

and convex posterior half suddenly narrows anteriorly

and, through the short ill-defined neck, passes to the relatively

small, narrow, obviously elongated head.

From

the ear region, the

head, continuing the general outline of the body, the

wedge

pro-

ceeds to the noticeably elongated, narrow facial portion with the somewhat extended, quite movable nose.

Limbs

short and massive, plantigrade with

naked lower surface

of feet. Claws strong, elongated (on middle digit of fore limb, 30-

35 along curve, 22-26 in straight line) and obtuse at end, adapted to digging. Tail short, approximately equal to head length.

mm

mm

Ear pinna relatively very small, rounded. Bare

large, separated

tip

of nose quite
fur.

from mouth margin by narrow band of

Eyes

are small.

The animal usually moves slowly and heavily, as if listless, its head. The posterior part of its trunk appears higher than the shoulder and the general appearance is depressed. The
lowering

badger does not flex the back as martens, polecats and wolverine do, and does not stand erect like the honeybadger. However, it can
run quickly
at a gallop.

In winter fur, pelage

on back and sides

is

long and coarse,

consisting of coarse, even bristly guard hairs and sparse quite soft

undercoat.
is

The venter

is

covered with short, sparse hairs and skin

usually visible in the inguinal region. Length of guard hairs on

is

the middle of the back in winter

821

75-80 mm.
in its winter fur coat,
i.e.

Color of the Middle Russian badger

late in

autumn before

retiring for the winter is as follows. Throat,

lower neck, chest and legs black, venter black with light brownish

1235

820

Fig. 297. Badger,

Meles metes

L. Sketch

by A.N. Komarov.

tints,

inguinal regions brownish-gray. General color tone of

sides
is

dorsum and

a beautiful

and pure light silvery-gray, as

if

coated with translucent black ripples.

On

the sides, there are straw-

colored highlights, more in the lower part. Such coloration is determined by differently colored guard hairs which have a broad

(35-38 mm) dirty-whitish or straw-whitish basal part, and after it a wide (22-25 mm) deep black band and a white or slightly grayishwhite tip 10-12 mm long. On the sides, bases of hairs have more intensive straw tone, the black band is somewhat narrower, and


1236
white'. The tail is covered by long coarse and generally of the same color as the back. The head is white, with white covering lower lips and chin also, where it is sharply bordered by the black throat. Two pure black bands pass along the head, sharply differentiated from the white background. Each of these begins on the upper lip at the mouth opening anterior to the corner of the mouth, and initially passes directly upwards. Somewhat below the level of the eye, the bands pass backwards at a right angle and, rising somewhat upwards and getting wider, pass through the eyes and, getting still wider, cover the whole base of the ear and even a part of the parietal region. In this region, its width is greatest. Behind the ear, the band, sometimes widening somewhat again, extends along the dorsolateral part of the neck sometimes to its middle (sometimes noticed somewhat farther back) and here disappears, gradually merging with the color of the upper body. Sometimes, it slants downwards between the color of the side of the neck (corresponds with back color) and the white field below. The width of the band anteriorly is about 15 mm, and in the ear region

ends of the hairs

is

hairs, quite fluffy,

45-55 mm. The black tone of the band is more intense in front of the eye, between eye and ear, around the ear and directly behind it. In the anterior section, the band frequently has a dirty or grayish tone and its outline is not sharp. Most frequently, the initial vertical portion of the band is weakly defined or almost undefined. The black facial bands outline a wide white band with almost parallel sides which extends from the nose tip through the [midline of the] forehead and crown. This band covers the occiput and the anterior part of the neck and sometimes almost the whole neck, gradually merging with the color of the upper body. Sometimes it does not lose its breadth and sometimes it tapers to a point and is
it is

quite well outlined.

The white

fields of the ventro-lateral surface of the

head ex-

tend backwards as a wide band of quite even width, between the

black color of the lower head and neck and the black head band.
Posteriorly, these [lateral] bands usually extend to the

tance as the upper [mid-dorsal] white band, and sometimes they occur notably on the greater part of the
'Sometimes
in

same dismore neck's length between

it

is

stated that along the badger's spine, gradually spreading out

is

posteriorly, runs a

brown band. This

a misundertstanding

such

a pattern is absent

our badgers.

1237
the gray color
ally

above and the black below. Posteriorly, they gradu-

merge with the color of the upper body. The ears are black with wide (to 17-18 mm) white edges along the upper margins. This edging is well marked against the black area of the facial band. Claws are dark-horn with yellowish cast.

Summer

fur

is

much

shorter, coarser
is

and sparser. Length of

venter,

guard hairs along the back

hairs are particularly sparse

more than 50 mm. On the

in places, the skin is visible
is

and

through

them. Color of badgers in

summer

darker and more dirty dorsally,

with yellowish or ocherous tinge; black tones ventrally are not as deep, but brownish. Yellowish tinge of the fur is determined by the

corresponding color of the guard hair

822

tips.

In the badger, there

is

New

hairs attain their full

one molt, extending the whole summer. growth only in hibernation. Therefore,

summer

fur is actually old being replaced and newly growing. General tone depends in part on wear and dirtiness of the fur. Individual variation in badger color is quite obvious. Equally

lighter
color.

with several predominantly moderate "gray" types, there occur and even very light animals generally of an almost silvery

More

frequently observed

is

more

or less significant dark-

ening of general color tone of the dorsum, or the appearance on it of yellowish tones, giving the whole skin clayey highlights. Changes
in intensity

of color of the black bands on the head occur, and also

the clarity of the white tone,

which may be darkened.

A brown
some

tint

may appear along

in

the black ventral fields. According to

data,

each locality two color types are present

in this species
It is

lighter

and darker.
is,

Mention of color dimorphism however, without foundation.

(Stroganov, 1962)

ordinary fluctuating variation.

sharp mutational changes, melanistic, albino and erythforms are known, and badgers of yellow color (Neal, 1948). Sexual differences in color are absent. Newborn animals are covered with short white fur; in slightly older animals, fur is lighter
ristic

Among

than in adults.
pelage.

By

the first hibernation,

young animals don

very
light.
it

adult

With increasing

age, white portions of the hair increase in

badgers, they lighten, and

may even become

it

Geographic variation

in color is considerable and,

lies in

appears

that relative to the described color types,

somewhat weak

lightening on one hand, and in strong darkening on the other. In

the latter case, color
the

may

attain a

brown

tone, while light fields of

visible.

head become so dark that

facial

bands become poorly

1238

The extent, form and location of black facial bands change also. In some forms, they do not cover the whole base of the ear, but pass below it, usually only touching the inner margin of the ear
conch base, and usually are narrower. The skull (Middle Russian badgers) is quite massive and heavy, noticeably elongated and, if crests are disregarded, quite even in
outline, not angular

and moderately high.

The
region.

line of the dorsal skull profile, if the sagittal crest is not

is

considered,

convex, with the highest point in the interorbital

this point, the line of the facial part
is

From

descends

at

an

obvious angle. Even steeper

the profile of the nasal opening.

Posterior from the interorbital constriction, the profile line forms

a very gentle arch, but descending

more strongly

in the occiput.

The braincase

is

moderately developed, not swollen and not elonis

gated, rounded-oval in outline, while the facial part of the skull

elongated and relatively narrow.

Zygomatic arches are narrow anteriorly and widely separated

posteriorly, with their greatest width at the level of the articular

condyles. Zygomatic width

is

considerably more than mastoid. The

well-

arches themselves are strong and massive, curved upwards in their

posterior parts.

The

postorbital constriction of the skull

is

developed, but short and not sharply defined, and the skull width
here
is

slightly less, equal or almost, equal to interorbital width. In

is

very old individuals, a fairly sharp intersection

lining the orbit, the dimensions of
Its

formed here.

Supraorbital processes are short, but well developed, sharply out-

which are

relatively very small.

greatest diameter constitutes slightly less than half the interor-

bital

width.
its

The

orbit

opens posteriorly, but bone borders more

behind the toothrow,

than half

circumstance.
palate, especially that part lying

The hard
is

of the molars
the anterior

long and narrow. Width of the palate between the inner margins is approximately 3.5 times less than its length from

824

edge of the sphenopalatine notch to the alveoli of the middle incisors. The interpterygoid groove limiting the palate posteriorly is short and wide. Its length is only slightly more than the distance between the ends of the hook-shaped processes. The
tympanic bullae are of irregularly angular (triangular) form, slightly swollen medially and moderately compressed (flattened) laterally. They do not contact the articular process. Mastoid processes are strongly developed and form massive outgrowths behind the

1239

823

Fig, 298. Skull of the badger,

Meles meles L.

auditory tubes which open independently of them. Paroccipital processes are short, triangular in form, their bases in contact with the
posterior parts of the auditory bullae.

The longitudinal diameter of

the nasal opening

is

greater than the transverse. Infraorbital

foramina are very large, irregular oval-rounded in outline. Their vertical diameter is equal to the diameter of the canine or greater

1240
than
its

it:

greatest diameter

is

approximately half the diameter of

the orbit.
Sagittal (arrow-shaped)

and occipital crests are strongly devel-

oped

in old

males, the height of the sagittal crest

may be

equal
its

to half the interorbital

width and even more. This crest reaches

greatest height in the middle of the parietals, arching

posteriorly,

downward

and

its

end overhangs the occipital crest and occipital

is

region in the form of a hook.

The lower jaw

relatively light

and elongated;

its

ramus

is

almost straight; jaw height in the middle part constitutes about

one-third the height in the region of the coronoid process. Posterior to the dental region, the

lower line of the jaw profile

is

rises

upwards quite abruptly. The angular process

coronoid, broad and high
process
is

thick and short, the

its

height above the level of the articular

at this

almost equal to

its

width

same

level.

The

posterior

region of the lower jaw has a low longitudinal crest ventrally.

Sexual dimorphism in the skull

dimensions,
is

is

insignificant and, besides

less

development of crests and generally somewhat "infantile" appearance of females. In young

manifested
in

somewhat

animals, the braincase

less

is

more swollen,
less,

postorbital constriction

marked

its

width not

but equal to or even more than

young animals, it is distinguishable), protuberances, etc. of skull are weakly defined, crests are not developed or only beginning to form. Complete development of
interorbital width (in the very

the skull

is,

apparently, attained in the second year, although fu-

sion of sutures begins earlier.

either
in

Geographic variation in the skull, not counting the size, is weakly manifested or almost absent. There is some change predominating form of the infraorbital foramen, in part (sculp-

turing) of the skull.

The upper
Along
is

carnassial tooth

is

relatively small, in plan, trianguits

lar outline (not pointed)

its

length approximately equal to

tall

width.

the outer side,

it

bears one

cutting cusp.
its

The

first

molar

strongly enlarged in dimensions

area

is

not less than the

second, and greater than the area of the carnassial.

in

form with

parallel outer

and inner

sides, the

It is rhomboid main diameter lying

approximately along the line of the toothrow;

greater [longer] than the outer.

its

inner side

is

The

tooth carries

some low

cusps,

from which the two main ones are located along its outer edge and one lies more or less in the middle of its cup-shaped crown.

1241

The lower

carnassial tooth

is

large, strongly pointed, with three

well developed apices in the anterior part and with an elongated flat and concave posterior part ("heel"); in length it exceeds the
anterior part.

The small premolars (one upper and two lower)

first

are

of the cutting type. The very small

lost

upper premolar
its

is

easily

and

is

frequently absent in adults,

first

alveolus

is

also not

evident.

The

lower premolar

may

also disappear and the

dental formula

I-C-P-M14 2 = 38
3
1

-3141

becomes

I-C-P-M14 = 36
3

3131

or even

I-C-P M = 34

13

or perhaps asymmetrical.
flattened as a result of

With aging,
wear of
its

the upper molars

apices.
825

may be more
premolar

Loss of the

first

in different parts of the

range occurs

The same is true with the lower premolar; in a portion of the races, it has two roots, and in another one. The OS penis is almost straight, with the dorsal side slightly concave or somewhat curved in an arc, and oval in cross section. At its tip, it is strongly widened in the form of a spoon and pierced
in different percentages of individuals (geographic variation).

in the

middle with a small foramen elongated

in the longitudinal

direction.
teristic

The sharp hook-shaped bend of

length
is

the anterior end characis

of the subfamily of martens, Mustelinae,

not formed.

600-900 mm,^ tail length 120-240 mm, length of hind foot, 75-130 mm, height of the ear is 35-70 mm. Condylobasal length of male skull is 113.0-144.0 mm, of fe-

Body

zygomatic width of males is 68.0-90.0 mm, of females, 62.0-85.5 mm; interorbital width of males is 25.2-34.5 mm, of females, 22.0-33.0 mm; postorbital width of
males
is

males, 100.0-140.0

mm;

19.9-27.5
is

of males

55.9-71.9

proximately
of]

mm, of females, 18.1-26.2 mm; mastoid width mm, of females, 52.3-67.0 mm (from ap300 males and females of the Z[oological] M[useum
Siberia).

M[oscow] U[niversity] and Stroganov, 1962, Os penis length is 72-76 mm.

^Body length of "about 90 cm" was stated by Ognev (1931)

as the average length

of old male badgers in

as maximal.

Moscow

province.

The majority of authors

refer to this length

European badger as 105 cm, while Kashchenko (1900, 1902) stated that the greatest length of the Transbaikal badger is more than 100 cm. These data are, apparently, somewhat exaggerated, although the length of 90 cm is possibly not maximum.
refers to greatest length of

Only Kuznetsov (1952)


1242

Weight of badgers at the time of their active life changes very growing from spring to autumn, and reaching its maximum entry into hibernation. The badger accumulates very much fat, and
greatly,
relative to
great.
its

general dimensions,
in

its

weight

is

disproportionally

Average weight

autumn of badgers

in the
is

European part

of the country, the northern Caucasus and Siberia

a

about 16-17 kg;

few large and particularly fattened animals reach 20-24 kg in weight; reports of badgers of 30 kg (Siberia; Stroganov, 1962) and 30 and 34 kg (Moscow and Ryazan districts; Ognev, 1931)
are not reliable.

Weight of the Middle European badger in summer is 7-13 kg, autumn 15-17 kg and as an exception 20 and perhaps even 25 kg (Gaffrey, 1961). Average weight of English male badgers is 12.2 kg, females somewhat less. The maximum established weight of
in

males

is

19 and 19.5 kg and of females

is

17.2 kg. There exist unveriin

fied reports of animals of 20.4

and 22.6 kg

weight (Neal, 1948).

Females are somewhat smaller and lighter than males. Dimensions and weight of badgers vary somewhat geographically. Although these changes are not very sharp, extreme forms (Middle Russian Amur, for instance) differ considerably (V.H.).

Systematic Position

Only species

in the genus.

Geographic Distribution
Europe, Near and Middle Asia, southern and middle zones of Siberia, the southern Far East, China,

and Central Asia.

Geographic Range

in the Soviet

Union

This constitutes a considerable part of the species range

the country.

its

north-

ern half and part of the western, and occupies the greater part of

In the northwest, the northern border of the range begins in

"the southwestern corner of Russian Laplandia" (Pleske,

here, apparently the following
is

1887):

understood as districts to the west

of Kandalaksha on the south, including

Kuusamo
lat.

(presently

Kusamo,
tat

in Finland).

This

is

apparently the most northernly habi-

of the species in Europe, lying at about 67 N.

is

On

the

Kola

827

Peninsula, the badger

absent, but to the south of Kandalaksha,

j5

IJJ

1244
it is encountered all over Karelia and in particular, recorded at Ruvozer (Marvin, 1951, 1959). Thence eastward to Arkhangel'sk, the southern coast of the White Sea forms the northern border of the range. On Solovets Islands the badger is absent, and it is not known whether it occupies the northern parts of Onezhsk

Peninsula.

On
the

the eastern coast of the

White Sea (Zimnii [winter]

coast),

northern border of the badger's range begins north of Arkhangel'sk, slightly above 65 N. lat. (Shil'dlog village or farther north). Thence,
it

passes eastwards along this latitude to Kuloi

Mezen' [still] on 65 N. lat. (Leshukonskoe village). Between Kuloi and Mezen', the border gives a sharp sag to the descending to Karpogar on the Pineg south a narrow extension [river]. From Leshukon on Mezen', the range border directs itself
and farther
to

to the northeast along

an irregular line and reaches the region of

the source of Pesha river, flowing into

is

Cheshsk

gulf. This [point]

only 50-70

km

south of the Arctic circle.

the southwest of Ust'-Tsyl'ma on the
left tributary

From
Pechora,

here, the range border turns sharply to the southeast and,

to

proceeding somewhat
it

goes to the middle course of the

of the

Pechora the Izhma river south (about 70-80 km) of Izhma city. Thence, rising somewhat to the north, the border passes to the[main] Pechora at 65 N. lat. (data of V.Ya. Parovshchikov for 1962). Farther, the border goes to the Urals, apparently along this same latitude (Narodnaya mountain) or a little to the south to Lyapinsk

Urals (Flerov, 1933).

It is

possible that, in the European part of the country, the

in
at

range of the badger extends Thus, a badger was caught

some

places even

more

to the north.

Kuloi near Kar'epol'e (50-60

km

more south of Mezen'

city;

Zhitkov, 1904) and near Mezun' city

(southern border of forest-tundra; Parovshchikov, 1959) and even

on the coast of Cheshsk gulf near the mouth of the Pesha

occurrence of
data require
this

(forest-

tundra; Parovshchikov, 1959). In these places, the character of the

animal

is

not entirely clear, and the available

more

precision.
is

However,

it

is

evident that this far

penetration to the north

connected with river valleys.

In western Siberia the badger

was recorded near Salekhard on

the

Ob'

at the

Arctic circle (Stroganov, 1962; the most northern

The border sinks sharply to the and passes through extreme upper reaches of the Pur and east the
point of occurrence in Siberia).

1245
(about 63 N.
It is

lat.;

Yanushevich and Blagoveshchenskii, 1952*).

possible that the actual northern border in the Trans-Ural lies

farther to the south

somewhat
(about 64;

Berezov on a little lower Yanushevich and Blagoveshchenskii, 1952**) and corin the

near

responds more closely to the situation of the border

Urals

and eastward from the lower Ob'. The Berezov upper Pur and Taz is nearer to the permafrost border. It is possible that its distribution to Salekhard is a distribution just along the Ob' valley. However, on the other hand the animal was noted at Khal'mersede
north of the Arctic circle (Leble, 1953)^.

From
N.
lat.

the region of

Taz sources, the border, going eastward,

at

descends to the south and crosses Yenisei, somewhere

about 61

At

this latitude, or a little north,

it

goes into the basin of

Podkamennaya Tunguska where the badger is met with at Baikit at the mouth of Chunya in Tunguska (appr. 61 35': N. Naumov, 1934) and a little eastward on the Mutorya river (Podarevskii,
entirely possible that the actual border of the range on bank of the Yenisei proceeds more to the north the animal was noted (apparently transient) at the mouth of the Taimura river in Nizhnaya [lower] Tunguska (N. Naumov, 1934; this same place is sometimes referred to as the mouth of the Vivi river both rivers flow into the Tunguska at nearly the same place).

1936).

It is

the right

828

Farther to the east, information on the limits of badger distri-

bution are very few. Apparently, the border

first

directs itself

towards a
tends
the

left

[bank] tributary of the Lena

the

Ichera, then ex-

somewhat northeastward (perhaps along the Lena valley) to mouth of the Vitim (about 59 30' N. lat.; Maak, 1851; D.

Ivanov) and even to a point about 150 km farther south of Suntar on the Vilyui (D. Ivanov, about 60 40' N. lat.). This place lies on
the

Lena or very close

to

it.

In the latter case

it is

it

is,

very likely,

always found as a transient since

village on the

very distant

to

Kudu-Kyuel'

Olekma

is

approximately 115

km

in a straight line

*"1958" in Russian original **"1953" in Russian original

'The border
of
its

Sci. Ed. Sci. Ed.

in

western Siberia

is

generally poorly clarified, and information

is,

in part contradictory.

Thus, the earlier data (Slovtsov, 1892; Chugunov, 1915) speaks distribution in "Surgut Territory" and not north of 62 N. lat. in Surgut region

on the Ob'. According to I. P. Laptev (1958), the northern border of the range, beginning at Berezov, embraces*** Kazym and passes somewhat to the south of the sources of Pur and Taz, reaching the Yenisei river at the mouth of Podkamennaya Tunguska. ***Misspelled "okhvatyvaet", instead of "obkhvatyvaet" in Russian original Sci. Ed.

1246

from its confluence with the Lena (D. Ivanov). Transgressions to the two above-mentioned places and in part to the mouth of the Vitim are very rare 3 cases during 40 years (D. Ivanov).

From
arching a

the
little

mouth of

the Vitim, the border curves, apparently

east, including the

towards the

Bodaibo region, and

turns back towards the southwest, apparently passing to the Kirenga

river (to the west of Baikal) without reaching the northern extremity

of Baikal and somewhere here,

it

reaches the shore of the Lake

(from data of V.V. Timofeev)"*.

In Trans-Baikaliya, the outline of the range

no

less

compliin

cated than to the west and north of [Lake] Baikal.

this

The border

region does not, judging from everything, extend so far to the

north. Starting, apparently,

on the shore of Baikal somewhere around

it

the

mouth of

the Barguzin [river],

passes along the eastern slope

to the south

of the Barguzin range on the northeast, crossing the Barguzin [river]

at

approximately 55 N.

lat.

and turns abruptly back

to

or southeast, including the Ikatsk range lying along the left

of the Barguzin and just

(tributary of the

fails

bank reach the sources of the Ok'

lat.).

Uda about

53" 30' N.

Thence, the border bends sharply to the east and northeast, crosses the Vitim [river], proceeds through the region of the Amalat headwater a tributary of the Vitim (in its bend) and extending'

northeastward, passes somewhere south of the Kalar

including the sources of the i.e., nearly reaches 56 N.
plateau, or at least a part of

in all events

Olekma and
lat.

its

tributary, the Tungir;

(V.V. Timofeev, S. Anashkin).

Therefore, apparently, the range beyond Baikal envelops the Vitim

it.

above description, it follows that that part of the range located in the upper Lena north of Baikal (Bodaibo) is not directly united with Trans-Baikal. Not excluded is the possibility
the
that such a

From

complex picture of the northern range border in the Baikal region is formed on account of a deficiency of accurate data. However, it is most probable that this is the actual situation, and it is explained by the fact that the badger bypasses to the north and south of the northern Baikal and Stanovoi highlands, the southern and northern Muisk ranges and several ridges connected with them (Kodar, Udokan).
"On
the

map

of

Stroganov

(1962,

p.

162),

the

occurrence

of badger
is,

northeast of Baikit on the

apparently,

an

incorrectly

Podkamennaya Tungaska was mentioned. This plotted habitat on the Mutorga river.

1247
In Yakutiya, the

badger

of the fauna.

It

is

is apparently not a permanent member known only from previously mentioned

transgressions to the

and

mouth of the Vitim south of Suntar village, Kudu-Kyuel' village on the Olekma river. From that place on the upper Olekma in Trans-Baikaliya at
at
lat.

about 56 N.

the border goes southeast, south of the Gilyui

(Gassovskii, 1921) approximately through Solov'evsk to Zeya city

and then
scends

to the

middle course of the Selemdzha. Thence,

it

de-

more steeply to the southeast to upper Burei, a little below the mouth of the Niman and in the source of a tributary of the Burei the Tyrma. Farther along the Evoronsk lowland the
still

border ascends towards the northeast, to the middle course of the

Amgun', and then proceeds again

to the southeast, crossing the

Amur and
829

reaching the ocean

at

52 N.

lat.

(from materials of

A. Samsonov and V.P. Sysoev).

On
To
of the

Sakhalin, the badger

is

absent.

the south of the above-mentioned line in the

European part

USSR,

the badger

is

encountered to the state frontier and

beyond it. It is also all over Siberia. In Middle Asia, the badger is met with throughout the mountains and is only absent in individual localities, mainly in extreme high-montane regions. Thus, the animal is apparently absent in the high-montane Pamirs, most notably in the western [Pamirs]. Badgers do not, however, avoid deserts. In the Kyzyl-Kum, they are encountered at least along the edge;
they are also recorded in the Ust'yurt, at least the eastern [Ust'yurt]

(Bazhanov, 1951) and along the lower Amu-Dar'ya (Gladkov and

Nikol'skii, 1935).
In Turkmenia, the badger lives mainly in montane regions, occupying in particular the whole of the Kopet-Dag and Bol'shoi Balkhan and, apparently, the Gyaz'-Gyadyk mountains (left bank of upper Tedzhen), Kushka valley and mountains to the east of Kushka (Chengurek mountains), and over the expanse lying along the mountains (V.G. Heptner). In the Karakum, the badger is rare, but is, apparently, met with almost everywhere (Yu.F. Sapozhenkov).

There are some unclear points and contradictions

in the char-

acteristics of the northern border of the range sketched above,

and

compared

to several other

works,

may be

explained not only by

incomplete information and imprecision concerning transgressions

of animals, and by natural fluctuations of the border during severe

1248
conditions in the north of the continent.^ Apparently the badger,
like
last

some other

species, has

expanded

its

range northward in the

decade. Thus, according to relatively recent information

in the European part of from Arkhangel'sk to Karpogory on the Pinega, thence to the upper Mezen' (Koslan) and across the middle course of the Izhma to the right tributaries of middle Pechora the

(Parovshchikov, 1959), the northern border

the country shifted

Shchuger* and Podcher'e. The border therefore passed considerably to the south of 65 N.
lat.

Geographic Range outside the Soviet Union

The range occupies Europe,

Ireland,
is

to the

west including England and

and

to the south to the coast of the

Mediterranean Sea.

It

on the Balearic Islands, Corsica, Sardinia and Sicily, as well as

on the islands of the Aegean Sea except Rhodes, and, apparently, on Cyprus, the badger is absent; it exists in Crete. The northern border of the range in Europe begins on the Atlantic coast of

Norway
N.
lat.)

at Trondheim Fjord (about 63 30' N. lat.; Ekman, 1922) and, describing an arc, curves northward (to approximately 64 30'

reaching the Baltic coast about the latitude 63 N.

lat.

In

Finland, the border begins on the Gulf of Bothnia approximately at Raahe (about 64 40'), and beyond forms a bend to the south,

but in the east of the country again rises somewhat northward and nearly reaches 66 N. lat. (perhaps even as far as Kusamo; see
above).
In Asia, the range occupies all of Asia Minor, Syria

and Pal-

estine, Iraq (apparently, only northern), northern, northwestern

and

western Iran (except the extreme south and the belt adjacent to
southern Iraq) and northern Afghanistan.
It

may

penetrate, per-

haps, the territories connected with the Hindu Kush, to the

more

southern parts of the country contiguous with Baluchistan. In the

east,

the range extends to Kashgariya and Dzhungariya (details

unknown

it

is

apparently absent in deserts); the entire northern

half of the
lian Altai,

Mongolian Republic, southward including the MongoKhangai, Kentei, and sections lying to the south of these
extreme east of
it.

montane

parts of the country, the

In China, the

range occupies the entire western part of the country

from former
late springs, the

'In Arkhangel'sk district, after cold winters with much snow and number of badger falls sharply (V.Ya. Parovshchikov). Misspelled "Shchugor" in Russian original Sci. Ed.

1249

Manchuria to the extreme south. The range apparently extends to north of Hanoi. In the east, the the extreme north of Vietnam range in China everywhere reaches the Pacific Ocean or to the

borders of our country;

its

distribution to the west

is

not clear.

It

occupies Inner Mongolia, Gansu, Shanxi, Sichuan and, apparently,

Yunnan
north of
latitudes

or a part of this province (the opinion that badger are

little basis),
its

absent from this region has

and possibly, the extreme

Burma

(information of

occurrence in Arakan between

20 and 15 N. is erroneous). According to some data, the range also includes Tibet. In actuality, it does not, apparently, occupy all of this territory, which is less suitable for the badger's life, and occupies only a part of perhaps the southern and southeastern margins. Thus, the Chiit nese race was noted in Lhasa and Dzhangdze (Gyantse), southwest of Lhasa, and the plains north of Sikkim (Pocock, 1941). Perhaps the "Tibet" range extends through southwest China and Upper Burma as a projection eastward along the Brahmaputra valley and

830 regions

connected with it possessing relatively more favorable The range does not penetrate adjacent India (or Nepal) but the possibility of penetration of badger from the northwest southward to Chitral, and from the north to Baluchistan is not excluded. The range area also includes Japanese Islands^ and Korean
natural conditions.

Peninsula (V.G.).

Geographic Variation
Racial variation in the badger
is

considerable.

It

manifests itself

both in amplitude of change in several characteristics and in the number of existing forms. About 40 names have been given to the

badger and 24 subspecies are usually accepted, of which 11 are within our country (Ognev, 1931; Ellermann and Morrison-Scott, 1951; Stroganov, 1962). This number is evidently too great. Together with that, those attempts to revise the racial composition of
the species that have been

done do not reflect the actual situation and themselves represent an extreme in the opposite sense 4 forms for our territory (Novikov, 1956) and even 4 races for the whole

species (V. Petrov, 1953).

*It is

sometimes claimed
is

that

it

occurs on Hainan (Pocock, 1941; Tate, 1947).

last

This error

apparently based on a misunderstanding. Pocock (1941) offered data of

Delacour about its occurrence "in the foothills north of Hainan". The misprint for "Hanoi". This text was so used above.

word

is

1250

830

Fig. 300. Head pattern of various race groups of badgers above, European badger, group meles; middle "sand" or Siberian Chinese, group arenarius leptorhynchus; below, Far Eastern, group amurensis anakuma. Sketch by N.N. Kondakov.

1251

Geographic variation of badgers reveals itself in general dimensions, skull measurements (for the most part insignificant), form of the infraorbital foramen (a character not very well defined and permanent), form of the upper molar its relative width (as well), structure of roots of the second lower premolar, relative frequency of loss of first premolars, general color tone and in form of head pattern. Color characteristics are most stable, typical and distinct. There are some geographic peculiarities in individual details

of skull structure.

According

to color type, all races of

badger are clearly divis-

ible into three groups.

The remaining

characteristics

more or

less

agree with coloration. Races directly "intermediate" are absent;

however, several other characters create "transitional" features

separate populations.

in

1. Group meles, European badgers. General color quite light and pure gray, "silvery"; sides of body a little lighter than middle of back. A wide black, rarely black-brown, stripe passes through eyes and envelops ear above and below (entire base of ear pinna).

pure white stripe passes from nose between [black] bands across

forehead, behind ears and along neck. Cheeks and sides of head
white, the white stripe passing along sides of neck^.
relatively
831

wide

Upper molar
second lower

ratio of length to width less than 1:5;

first

premolar usually with two well-developed roots,

upper and lower

premolars for the most part retained. Infraorbital foramen usually more
elongated in vertical direction. Dimensions large to moderate.
In Europe eastward to the Volga, Caucasus, Near East, south and southeast Middle Asia.

Group arenarius-leptorhynchus, Siberian or "sand" badgsome forms, color corresponds to that of preceding group, and sometimes even darker, smudged with
2.

ers^ Color lighter, but in

ocherous and brownish highlights. Sides a

little

lighter than

middle

of back. Dark facial stripe usually not black, but brownish in tone;
it

posteriorly,

narrows behind eye and extends above ear. It does not go far sometimes hardly reaching ear. White color on head
^This type of coloration was described above in

more

detail in the section

"Description".

^Animals of

this type

Ryn sands between

the lower

were recorded in our country for the first time from the Volga and Ural [rivers]. The form described from there

by Satunin was called arenarius; i.e., sand. This Russian name was introduced in our literature for the entire group of races, although the majority of them have no relationship to sands (montane, forest).


1252

usually dirtyish. Light stripe passing along head dorsally between

dark stripes relatively narrow and short; it does not extend behind ear or only extends behind it for a small distance and already

merges with color of dorsal body on occiput. Upper molar relatively elongated ratio of length to width 1.5 or nearly that; second lower premolar usually with one root; first lower premolars for the most part absent. Infraorbital foramen more extended in horizontal direction. Dimensions moderate. In Trans- Volga; Siberia except Amur region and Ussuri basin; Middle Asia except extreme south and southeast; and all remaining areas of Asia except Near East and notheastern China (former

Manchuria) and Japanese islands. From a zoogeographic and systematic point of view, it is noteworthy that in Europe, the Volga, and in part, the Kama (lower course) rivers separate two sharply differentiated race groups

European and sand badgers. They evidently play the

role of an

insuperable mechanical [physical]* barrier for badgers

animals
(see

which

in these latitudes are not active in winter. This fact has

multiple significance both in several general respects (problem of

barriers,

problem of

clines,

and others).

If in

some forms

below), individual "transitional" characters are noticed, they do not apply to facial pattern. At the same time, the limits of distribution of both groups of badgers in the northeastern European part of the country, and in the northern Ural region are not clear. In the

Pechora-Ilych preserve (upper Pechora), the Siberian badger exists. In Tien-Shan (see below), the races of both groups also approach each other closely, but are sharply separated by mountain
ranges.
3.

color very dark, with

Group amurensis-anakwna. Far Eastern badgers. General brown tones strongly developed. Head pat-

tern corresponds to that of the Siberian badgers; however, head

dark, and dark stripes weakly defined.

Upper molar
root, first

is

elongated, second lower premolar with one

premolars always absent. Infraorbital foramen wider in

region, Ussuri basin, northeastern China (former

transverse direction. Dimensions small.

In

Amur

Man-

churia), and Japan.

Badgers of groups 1 and 2 were, up to the 30' s, considered M. meles and M. leptorhynchus (Ognev, 1931). different species
*In Russian original, the

word mekhanicheskoi

is

used

Sci. Ed.

1253

The Amur-Ussuri badgers were

(V. Petrov, 1953),

affiliated with the latter.

The

recent attempt to restore this completely neglected point of view

was naturally unsuccessful. scheme of races given below is preliminary. The

832

Group of European badgers, meles.

1.

Middle Russian badger, M. m. meles Linnaeus, 1758

(syn.

tauricus, caucasicus).

Dimensions large. Color of back relatively pure silvery-gray tone. Main tone of head pure white, dark stripes wide, black in color, white fields extend far backward along upper and lateral parts of neck.^ Pelage
relatively soft with relatively dense underfur.

Skull with strongly developed crests, second lower premolar

with two separate or merely adjacent roots;

all,

first

premolars, often
ratio of length to

usually present; upper molar relatively wide

width about 1.25, and usually not more than 1.35. Infraorbital foramen extended in vertical direction, and its greatest diameter, being directing upwards and somewhat internally, greater than transverse diameter.

Dimensions maximmal for its group; apparently, for entire speMiddle Russian badgers are largest they are, evidently, larger than Middle and Western European. Condylobasal length of male skull is 125-140 mm, of females, 123.2-133.6 mm; zygomatic width of males is 75.2-89.5 mm, of
cies.

females, 68.0-85.2

mm; mastoid width of males is 62.1-71.9 mm, of females, 58.5-67.0 mm; length of upper molar of males is 14.717.1 mm, of females, 14.5-17.1 mm; greatest width of posterior molar of males is 1 1.2-13.2 mm, of females, 1 1.0-13.5 mm (Ognev,
1931;

some deviation

in

both directions possible). In animals of

this race, different

parts of range

south

60%,

in

numbers of first premolars retained in different in Middle and Northern Russia, only 77%, in the Cis-Caucasus 66%, and in Crimea about 72%

(V.G. Heptner).
a little

Weight up to 20-24 kg (autumn), in exceptional cases, perhaps more (see above). In European part of Union eastward to Volga, in the Crimea,
to east in north

Cis-Caucasus and northern Caucasus. Distribution

^The above-given description (page 1234) applies
to this

form.

1254
of European part of country not

known

(see below, separation

from Siberian badgers).

Outside the

USSR,

in

remaining parts of Europe except Rhodes,

Crete, Spain (existence of a separate race in

Denmark

is

unlikely).

Badgers from the northern slopes of the Caucasus, its foothills and adjacent plains may be, on average (skull dimensions) slightly smaller than Middle Russian. In this one might see some tendency
toward transition to the Trans-Caucasian form. This difference is, however, completely insignificant and separation of a different form,
caucasicus,
2.
is

not justified.

Kizlyar badger, M. m. heptneri Ognev, 1931.

stripes

Dimensions large (as in the preceding form). Color very pale, dull, dirty-grayish-ocherous. Black head narrow, but typical of European group.
Skull as in preceding form.

on

In Caspian steppes area (steppes of northeastern Cis-Caucasus), right

bank of lower Volga (Kalmytsk steppes), and Volga

delta.

Outside the

USSR absent.
known form;
is

very poorly

exhibiting several characters simi-

lar to the

Siberian badger group, but on the whole typical of Eu-

ropean [group]. Range

very small, especially as compared to

other subspecies, but on the whole, characteristic of a series of mammalian races, mainly of eastern origin, which are also restricted in their distribution to the above-mentioned section of steppes,
deserts and semideserts.

This form requires further study and here

ognized.
833

is

provisionally rec-

As

yet there

is

no basis for placing

it

in

synonymy with

the

nominal form. 3. Trans-Caucasian badger, M. m. canescens Blanford, 1875

(syn. minor).

Dimensions smaller than preceding forms.

brown

General color of dorsal side dirtyish-gray with highlights of tones. Head color as in nominal form. Skull and tooth structure as in nominal form; i.e., infraorbital foramen high, but upper

molar somewhat elongated, though not so strongly as in Siberian badgers, and crests somewhat more weakly developed. Condylobasal length of male skull is 116.0-123.3 mm, of females, 111.8-122.0 mm; zygomatic width of males is 68.0-81.5

1255

mm,
63.2

of females, 66.0-72.0

mm;

mastoid width of males

is

56.0-

mm,
is

of females, 56.1-58.0
its

mm;

length of upper molar in

males

15.9-17.3,

width 10.0-11.1 (Ognev, 1921). In badgers

of possible

of this form, about

57%

number of

first

premolars are

retained (V.G. Heptner).

In Trans-Caucasus, Kopet-Dag and apparently, Gyaz'-Gyadyk mountains (right bank of upper Tedzhen); probably elevated regions of Kushka (Chengurek mountains) in Turkmenia. Limits of

distribution in northern

Turkmenia not known.

Iran, Afghanistan,

Outside the

USSR in

and possibly Asia

Minor.

Fergana badger, M. m. severzovi Heptner, 1940. Dimensions quite small, apparently close to preceding form. Color of dorsum relatively pure silvery-gray without yellow sheen or with only insignificant development of it. Black stripes on head wide occupying whole ear. Upper molar short and broad; i.e., typical of European badger group, preorbital foramen extended in transverse direction; i.e., a
4.

character of Siberian badgers.

Condylobasal length of skull (10) is 109.4-M 118.4-127.0 width is 64.5-M73.2-81.6 mm. In animals of this about 26% of the race, the least number of the first premolars

mm; zygomatic

number

possible,

among

all

groups of races (V.G. Heptner).

In region of right tributaries of

Pyandzh and upper Amu-Dar'ya,

it

Pamiro-Alaisk system (apparently. Eastern Pamir only; the Alaisk

valley),

Fergana valley and mountains bordering

on south and

north (Chatkal and Fergana ranges).

Outside the
ever,

USSR absent.
closest to

The Fergana badger stands

it

M. m. canescens.* Howseveral skull characteris-

is

substantially different

from

it,

tics "transitional" to the

Siberian badger group in particular being

most noticeable. It is probable that its range is separated from that of M. m. canescens by the Pyandzh [river]. There is a remarkably sharp boundary between its range in the north and badgers of the leptorhynchus group (sand) which occupy the northern ranges of the Tien-Shan system. Already along the southern slopes of the Talas Alatau south of Issyk-Kul, and along the upper Naryn are distributed typical representatives of the Siberian badger group.

The

limits

between both forms

in the plains are not clear. In all

*See above, Trans-Caucasian badger

Sci. Ed.

1256
events,

. .

severzovi was recorded in Kugitangtau (Ishunin,

1948).

1961), and M. m. leptorhynchus in Nuratau (Kuznetsov,

partially contradictory (Kuznetsov, 1948; Ishunin, 1961).

Other data on badgers of Middle Asia are poorly defined, and

Group of

Siberian, or "sand" badgers

arenarius

leptorhynchus

5.

Siberian badger, M. m. sibiricus Kastschenko, 1900 (syn.

altaicus, raddei, aberrans).

Dimensions moderate
834

somewhat smaller than

in

nominal form,

but larger than in Trans-Caucasian.

General color tone of back light

typical for

gray, usually with yellowish

Dark stripes on head and extend along dorsum of head, not including ear. Their color varies from brownish-black to tawnybrown. Pelage long and relatively soft with dense undercoat.
or straw-colored and sandy-straw highlights.

group

narrow

Upper molar
1.5.

relatively elongated, ratio of

its

length to width about

Infraorbital

foramen extended
root,

in

transverse direction, second

lower premolar with one

and

first

premolars very rarely occur.

Body length of males (6) is 675-M720-750 mm, of females 620-M668-692 mm; tail length of males is 185-M214-235 mm, of females, 174-M 196-222 mm; length of hind foot of males is 100-Ml 16-127 mm; of females, 82.0, M84.2-86.0 mm; ear length in males is 38-M39-41 mm, of females, 37-M38-40 mm.
(4),

(7) is 123.0-M125. 8-128.7 17.4-1 18.7 mm; zygomatic width of 116.0-Ml mm, of females, (5) males is 75.6-M78.0-80.5 mm, of females, 68.4-M69.2-70.2 mm; interorbital width of males is 27. 3-M27. 8-28.3 mm, of females, 26.2, M26. 6-27.0 mm; postorbital width of males is 22.0M23. 8-24.6 mm, of females, 20.0-M23.2-24.0 mm; mastoid width

Condylobasal length of male skull

of males

is

64.0-M68.7-75.4

mm,
first

of females, 58.6-M60.2-62.0

mm.

Percentage of retained
populations of this race

premolars differs in different

in

Bashkiria and southern Urals

in

about

13[%], in Siberia and Altai 3[%] and (V.G. Heptner).

Trans-Baikaliya 0[%]

Weight of adult males reaches 10-13.6 kg, and evidently even more (all data on dimensions after Stroganov, 1962).
In Siberia including Trans-Baikaliya

and

Altai, northern parts

of Kazakhstan and probably northern Trans-Volga.

1257
Outside the USSR unknown. The independence of the Siberian form of badger was sometimes, and is now, doubted. At the same time, its differences from the form arenarius, and from the Mongolian form leptorhynchus,
is

sufficiently real. Separation of this race

is

arguable, but the

example, badgers from the middle course of the Ob' and from the deserts and semideserts of southern Kazakhstan and Uzbekistan one may hardly admit even theoretiracial identity of, for

badgers from the China and Tibet belong to one form (Petrov, 1953; Novikov, 1956) acceptable. The separation of the form aberrans Stvoganov, 1962, demands confirmation. This form is, to a high degree, doubtful, both as
cally.
is

Neither

the

view by which

all

Trans-Caucasus, Siberia and Turkmenia

to Central Asia,

its characters and its range ("northern regions of eastern Kazakhstan" Tselinograd, Kokchetav, Pavlodar, Zaisan Lake and middle Tarbagatai). Concerning Trans- Volga badgers, see the following form.

regards

6. Kazakhstan or sand badger, M. m. arenarius Satunin, 1895. Dimensions moderate somewhat less than in nominal form,

but greater than in Trans-Caucasian form.

Color is lighter and paler than in northern forms, conditioned by decrease in black bands of guard hairs and their frequent full reduction on sides of body, where the color acquires a pale-straw
tone.

clayey tone usually develops over the entire skin, which

Pelage

impinges on the silvery-gray color characteristic of northern races. is coarse, bristly and underfur is scarce.
Structure of skull and dentition as in Siberian badger, M. m.

sibiricus.

835

700-780 mm, of females (4), 610200-220 mm, of females, 185-250 mm; length of hind foot in males is 115-126 mm, of females, 95110 mm; ear length in males is 58-70 mm, of females, 40-80 mm
length of males (4)
length of males
is

Body

700 mm;

tail

is

(Sludskii, 1953).

Condylobasal length of male skull is 115.0-131.0 mm, of mm; zygomatic width of males is 68.6-77.7, of females, 62.2-73.8 mm; mastoid width of males is 55.9-65.0 mm, of females, 52.3-58.8 mm; width of upper molar in males is
females, 109.0-118.0

10.1-12.1
in

mm,
first

of females, 10.1-12.3

mm;

length of upper molar

males

is

14.1-16.6

mm,

of females, 14.0-16.0

of retained

premolars

about two (V.G.

mm.

Percentage

Heptner).

1258

Weight of males in March-May is 7.8-8.3 kg, in March-June is 5.6-7 kg. Weight of young animals in November is 10.5-12.5 kg, of an adult female, 14.5 kg (Sludskii, 1953). In southern Trans- Volga (Volga-Ural steppe), Kazakhstan except the northern and montane parts, and plains of Middle Asia except the parts occupied by the forms canescens and severzovi (extreme southern Turkmenia, Pamir-Alaisk mountain system, Fergana valley and the mountain ridges bordering it on the north).
Outside the

USSR absent.

This

described
not clear.

form of badger
Its

is

morphologically

and

geographically well-characterized.

distribution in northern Trans-

Volga

is

It

was recorded

in the southern Urals (Kirikov,

1952), at the latitudes of Samarsk

Luka and Bashkiria. However,

its

these data indicate only the presence of badgers belonging to the

"sand" group

in these places,

but they do not confirm

identity

with the form arenarius from the Volga-Ural steppe and Middle Asia. This is all the more so, since badgers of northern Kazakhstan

belong to the Siberian form sibiricus.

It is

possible that these Trans-

Volga badgers (except southern) also belong there. In general, as was shown, limits of distribution of the sand badger group to the
north in the Cis-Urals are not

known

to

be contiguous with the

meles group.

form in the steppes of the Cis-Caucasus ("Caucasian steppe", Ellerman and MorrisonScott, 1951) was mistaken. As was shown, all of the forms of the leptorhynchus group are absent west of the Volga and Caspian
Information on the occurrence of
this

Sea.
7.

Tien-Shan badger, M. m. tianshanensis Huene, 1910 (syn.

talassicus).

Dimensions moderate. General color same as type, and

m. arenarius, but

as in

somewhat

darker, the yellow sheen

Kazakhstan badger, M. is weaker or

not developed, the fur longer, denser and fluffier. Percentage of

retained
first premolars is about 8.3% (including material from around Dzhungariya; V.G. Heptner). In the northern ranges of the Tien-Shan including Dzhungarsk,

Zailiisk, Kirghizsk

and Talassk [Alatau]. In the eastern parts of the [mountain] system within the USSR, southwards as far as the upper
[river].

Naryn

Outside the

USSR probably

in

the Tien-Shan system in

Dzhungariya and Kashgariya [China].

1259
completely unclear and entirely doubtful form which
It is

It

is

requires additional study and confirmation.

to the highest

degree close to the Kazakhstan badger, M. m. arenarius (many

individuals are not distinguishable at
all)
is

and apparently, differ

only in somewhat denser underfur, as

frequently the case with

montane populations. The southern limits of this form and the whole "sand" badger group requires more precision. The Fergana badger, M. m. severzovi
(see above), exists along the southern slopes of Chatkal and, prob-

Fergana ranges and Tien-Shan form along the upper Naryn. The problem of variation and nomenclature of badgers of the northern Tien-Shan is not clear. Hoiningen-Huene (1910) named the Tien-Shan form from living animals received from
ably,

Note.

Hagenbeck* and caught

"Tien-Shan" but not noted whether within the borders of the [present] USSR or in China. From phoin
it

tographs and the description

is

clear only that this

is

a typical

"sand" badger. Ognev (1931), without attempting

the type locality
836 cated to
it

to precisely limit

form tianschanensis, alloChinese parts Kaitsalai, Yuldus, Bororiver, (Kul'dzha, Kunges Tien-Shan of the distribution of Khoro range). At the same time, he described the this form, according to data of N.A. Severtsov, in a series of places in the eastern parts of the Tien-Shan within the borders of the USSR westward to the Chu river as well as in the western TienShan, "for the entire range in general" and even for the Karatau
(t.

t.** restricta) of the

individuals from the extreme eastern

(page 478). Factually, therefore,

it

is to

take the habitation of the

form

tianschanensis throughout the entire Tien-Shan, within the

borders of our country and beyond them.

On a par with this, in the same work [Severtsov, 1873] (pages 478-479), M. m. talassicus was described from Talassk Alatau (southern slope) from two specimens; the occurrence of which is assumed to be also on the Chu river (Frunze-"Pishpek"***). Differences from the form tianschanensis given are very insignificant
sis

(somewhat darker color). The place of origin has not yet been accurately established
tirely

for

doubtful

and besides

M. m. tianschanenit

is

en-

that

it

is

possible that there are different badgers

of the type arenarius-leptorhynchus from the different parts of

Tien-Shan, there

is

no basis for admitting the independence of the

form talassicus. German animal

dealers

Sci. Ed.

**For "terra typica";

i.e.,

type locality

***Presently Bishkek, Kyrgyzstan

Sci. Ed.

Sci. Ed.

1260

Group of Far Eastern badgers, amurensis-anakuma.

8.

Amur

badger, M.

m.

amurensis Schrenk,
in the

1858 (syn.

schrenkii).

Dimensions very small, smaller than

badger M. m. canescens.
It

Trans-Caucasian

is

the smallest race of the species.

all

General color very dark, darkest among

races.

Dark

stripes

on head extend above ear and disappear in dark field of occiput; they are black or dark blackish-brown or sometimes coal-black in color. Entire area between stripes and cheek region, i.e., places

which

in

other races are white or very light, are dirty grayishcolor.

It

brown

in

may be

so dark that stripes

are

weakly

distinguished or almost undetectable. Color of back very dark due

development of dark bands of the hairs, grayish-brown with silvery highlights, depending on white tips of hairs. Sides of body only slightly darker than back. Pelage with little wool, but soft.
to strong

Skull small, with smooth outlines, relatively light. Protuberances, etc. on skull weakly defined, and crests very weakly

developed even
tended
in

in oldest individuals

(wear on crown of carnassial

teeth). Entire skull has infantile features. Infraorbital

foramen ex-

transverse direction, upper molar

is

elongated. First

premolars always absent (extreme degree of reduction within

limits of species).

Body length 60-70 cm. Condylobasal length of male skull (6) is 111.0-117.9 mm, of females (6), 104.8-112.2 mm; zygomatic width of males is 67.875.4 mm, of females, 62.1-73.2 mm; mastoid width of males is
57.3-63.5

mm;

of females, 53.0-58.2

mm;

interorbital width of

males

is

24.0-29.2

mm,

of females, 22.1-24.6

mm;

postorbital

width of males is 20.8-22.4 mm, of females, 18.2-21.9 (Stroganov, 1962 and materials of Z[oological] M[useum of]

mm

M[oscow]
17.0

U[niversity]). Length of upper molar in males

is

16.2-

mm,

of females,

14.0-15.3

mm;

width of upper molar in

(Ognev, 1931). males is 11.2-11.5 mm, of females, 9.6-10.8 Badgers of this form are always characterized by complete
absence of
first

mm

premolars and corresponding change in dental for-

mula (V.G. Heptner).

In Ussuri Territory and Priamur'e westward approximately to 123 E. long. (Albazin).

1261

Outside the
shoi [Great]

USSR in

contiguous parts of northeastern China

(former Manchuria), and probably westward to include the

Khingan [range] and Korean peninsula.

is

The Amur badger

pattern,
it

a sharply distinguished form. In

is

its

head
satufirst

resembles races of the "sand" badger group, but

all

sharply
its

separated from them and

other forms of the species by

rated darker color, by size and complete reduction of

all

by dental formula. This form, undoubtedly, possesses features similar to the Japanese badger, M. m. anakuma; however, the union of both races is unfounded.
premolars;
i.e.,

837

The borders of
fully clear.

the range of the

Amur

subspecies are

still

not

There was some information

that badgers of Trans-

Baikaliya possess characters transitional to the

this
is

Amur

form, but

not so. In the Mongolian Republic, near Beijing, and

possibly in the central Manchurian plains lives a badger of the

typical "sand" type, extremely light.

The Amur badger

istic

is

a clear

example of one of the character-

features of geographic variation of

species of Palearctic
color,

mammals

darkening

some widely distributed or more saturation of

and partly decrease in size in the south of the Far East {Cervus elaphus, Capreolus capreolus* Mustela sibirica and
several others).

Beyond
majority, of

the borders of our country, a quite large

number of
if

farms have been described. The independence of many,

not the

them needs confirmation. Usually accepted are: 1) M. m. anakuma Temminck, 1844 Japanese Islands (Hondo [Honshu], Shikoku, Kyushu, ? Hokkaido); 2) M. m. melanogenys J. Allen, 1913 Korean Peninsula (evidently a synonym of M. m. amurensis southeastern Tibet V.H.); 3) M. m. leucurus Hodgson, 1847 China, (Lhasa); 4) M. m. leptorhynchus Milne-Edwards, 1867 Mongolian Republic'"; 5) M. m. blanfordi Matschie, 1907

pygargus Sci. Ed. considered a distant species, '"Osgood (1932) considers the forms leucurus and leptorhynchus identical. This point of view was, apparently, also supported by G. Allen (1938). In this case, the Chinese-Mongolian badger must be called leucurus (not leptorhynchus as accepted by Allen). In general, the identity of badgers from Lhasa and from around Beijing (the
type localities of both forms)
is

*Now

very doubtful, and this question needs further study.

to leucurus

Allocation to leucurus of the badgers of Siberia, Middle Asia and even Trans-

Volga (Bashkiria; V. Petrov, 1953) belong

is,

of course, not accepted.


1262

. . marianensis Graells, 1897 Pyrenese Peninsula; 7) M. m. arcalus Miller, 1907 Crete; M. m. rhodius Festa, 1814 Island of Rhodes, Aegean Sea; M. m. ponticus Blackler, 1916 Asia Minor (very probably, a synonym
Kashgariya; 6)
[Iberian]
8)

9)

of M. m. canescens; V.H.); 10) M. m. danicus Holton, [1935]

Denmark

(evidently a

synonym of

the nominal form) (V.H.).

Biology
Population. Badger populations within the limits of
variable.
It is

its

range are

determined chiefly by the abundance of food, breed-

ing burrow conditions and sources of drinking water. In the breed-

ing period, in the large spruce swamps, and on the water divides

of the Volga and Zapadnaya [western] Dvina,

great, reaching

its

density was not

in is-

0.13-0.16 per 10 km^. In the forest-steppe,

land of broad-leaved oakgroves (Tula abatis; Likhachev, 1956),

average density over 13 years was from 1.6 to 2.1 per 10

km^

In

838

one of the great islands (28 km^) of the delta of Hi river in Kazakhstan (Sludskii, 1953), density reached a maximum of 21.026.0 per 10 km^ (15-16 litters). Data on the number of inhabited burrows are less reliable: some burrows are only periodically inhabited and in others, up to three litters live (badger "city"). In the European part of the USSR, the badger is most numerous in the middle zone there where islands of forests prevail, and the locality itself is hilly and cut with ravines. Thus, in Tatariya, 3-9 animals are found in 10 km^ of wooded lands suitable for the habitation of badger (Gorshkov, 1964). The greatest number of badger skins are obtained from Kazakhstan, although the animal itself is far from numerous throughout the region. The least number of badgers is in the northern Kazakhstan steppes. It is most numerous in Alma-Altinsk, Dzhambul and Chimkent districts (Sludskii, 1953). In the taiga, both lowland and montane, the badger is rare. The same must be said for the open steppes. Habitat. The badger reveals great flexibility with respect to habitats. Just as in steppe and forest foxes, wolves, tundra and taiga reindeer, etc. badgers of various races differ in their ecology. The badger in the forest zone is a forest animal, mainly restricting itself to forest edges, forest islands, coppices and wooded ravines. It is most numerous in the zone of islands of mixed forests

1263
alternating with fields,

meadows and
food for them.

settlements.
if

It

avoids large

forest massifs, rarely settling those,

and

so mostly along edges;

the contrary, in a small

such forests have

badger. In

little

On

island forest of 15-20 h, there are sometimes up to three litters of

some

localities (Byelorussia,

Kirov

district), the

popu-

lation of badgers corresponds to the percentage of

woodlands, which

is explained by the excessive pursuit of the badger by man, and not by environmental conditions. In Moldavia (Korchmar', 1963), the badger inhabits the forest tracts of Kodra, in insular forests of the

northern regions and in southern forests.

It

is

small islands of forest. For the badger, the forest

most numerous in is merely a good

refuge sheltering

it

during foraging, and not a necessary sur-

rounding for

its

life.

In the habitat, besides

abundance of food, favorable burrowing

is

conditions are the most important requirement. This

natural for

an animal pursuing a semisubterranean

spent in the burrow.

way

greater part of the [24 hour] day and several winter

The badger prefers dry, deep ground water level for easily-yielding excavation. In extreme cases, it settles in more or less moist soils, choosing the most elevated, well-drained places. Forest attracts the badger not
a

which the months are sandy-loam soils with

of
life, in

only as a shelter, but also thanks to the presence of the Ortstein

horizon which
is

impermeable

to

water (Kolosov, 1935), under

it

which

it

digs

its

burrows. For the same reason,

as the vault of

prefers sand

lying beneath a layer of clay or loam.

The water-impermeable upper

its

layer of soil then serves

it

underground construcin

tions (Lukhachev, 1956).

The badger sometimes digs burrows

sands of pine forests, and also

especially

among

rocks.

The second condition for the badger is the proximity of water, when the approach to it from the burrow is hidden. In

open dry steppes of southeastern Trans-Baikaliya, habitats of the badger are always connected with water resources. In Kazakhstan, the badger sometimes makes its burrow not more than 5-10 metres from water (Sludskii, 1955). The badger does not avoid human
settlements as long as the burrow itself
lated,
is

located in a little-popu-

hidden place, most often on the slopes of a deep ravine

overgrown with trees, bushes and tall grasses. In the middle zone, the badger prefers the edges of mixed
forests but
it

does not avoid coniferous forests.

And

here,

it

prefers

sections dissected by ravines, but sometimes settles on flat plains.

1264

839

Montane oak and mixed coniferous-broadleaved forests. Typical habitats of in southern Primor'e. "Kedrovaya Pad' " preserve. Here also exists a fauna complex of Ussurian Territory kharza, racoon dog, white-chested bear,
Fig. 301.

Amur

badger

leopard, and others. July 1964. Photograph by A.G. Pankrat'ev.

In the forest-steppe zone, the badger

more often burrows along

if

the

slopes of ravines and gorges which are very often covered,

not

by forest then by steppe shrubs and rank growth of grasses. In Kazakhstan, the badger is an animal of steppes and semideserts, ecologically different from the badger of the forest
zone. In northern Kazakhstan,
it

settles

along the slopes of ravines

1265

woods, on high and along the river valleys. In central Kazakhastan, it settles in the bank precipices and the slopes of hills and ravines, especially the shores of lakes and rivers. In the desert zone, it lives in consolidated sand hills alternating with solonchaks, streams and
(saev), the high ridges near lakes in islands of pine
terraces,

lakes, in flood lands and deltas of rivers. Many badgers are also found on "terraces"* of foothills, especially in apple forests on the

slopes of mountains. In mountains here and in the Caucasus

it it

ascends to a height of 2300-2500 m. In the montane Altai,

restricts itself to the

lower belt of the mountains

not higher than

dry valleys
at the

1000-1200 m.
In Ussuri Territory (Yu.A.

badger

is

associated with more gentle relief.

Salmin and V.D. Shamykin), the It is encountered in

in well-drained

valleys of large mountain rivers foot of slopes.

839

brown

Food. Of the whole order of carnivores the badger, like the bear, is the least carnivorous. In the badger as well, the
teeth

development of powerful
ten with

and masticatory musculature

is

con-

nected not with camivory, but with omnivory. While in the pine mar-

mixed feeding

the intestine length

it is

suasses body

4.5-5 times, in the badger

times).

8 times (in the bear,

length by by more than 10

The badger's food

ards), insects

consists of rodents (mainly voles), rarely

liz-

small birds, amphibians (mainly frogs) and reptiles (mainly

nuts,

and their larvae, molluscs, earthworms, various berries and other plant food.
it

fruits,

The species composition of food, and within

each food type,
840
is

the ratio of

subject to considerable variation, both seasonal

and geographic. The effect of geographic variation upon food

composition can be judged by Table 70, which presents the data

on the most important foods of 7 separate parts of the range.

Actually, the diversity of the food components
is still

more

signifi-

cant. Apparently, in the northern parts of the badger's range there

more camivory, less in the south where sources of its food are more abundant and varied (Crimea, Kazakhstan). Among mammals, common and red-backed voles are the most important. The role of birds is not great. Usually, they are small
is

birds nesting on the ground, mainly during the nesting period, or

some kinds of wounded

birds.

The shores of

large bodies of water

Sci. Ed.

*In Russian original "prilavok," or counter, shelf

1266

''

L.
840
Fig.

302. Broad-leaved forest in the valley of Kedrovaya river

typical habitat of

Amur badger "Kedrovaya

Pad'

"

preserve,

southern

Primor'e.

October

1963.

Photograph by A.G. Pankrat'ev.

constitute an exception (Kaletskaya, 1957). In destruction of eggs

and young of game birds, badgers, for the most

without cause. In the forest zone, frogs are,

part, are

accused
of great

in places,

importance. In the dry forests of Zhigula, the importance of

slow-worms [anguine
forest

lizards]* increases, while in the

Buzulak pine

it

is

lacertid.

Among
dung

invertebrates in forests, of greatest significance are

ground beetles, stag-beetles, longhorn beetles, forest may beetles and June beetles and their larvae, sawflies, wasps and bumblebees and their larvae. The badger eats many mollusks (mainly slugs) and earthworms (in large amounts). Among plant foods, of greatest importance are acorns, hazelnuts, strawberbeetles, various
ries,

wild apples, pears and others. In Kazakhstan, the main foods

of the badger consist of insects, mainly mole crickets, and in

years

some

Asiatic
Sci.

locust on

which
is

it

is

quickly fattened (Sludskii,

*The Russian word "veretenitsa"

laevis

applied to both Anguis fragilis and Coronella

Ed.

1267
841

Table 70. Geographic variation

in

badger foods (occurrence, %)

Buzuluk
pine
1

Food type

Tula Abatis Tataria (Likhachev and

,

Zhiguli
(P.B.

Montane Crimea
(Savina,

Delta of Ryabinsk
Hi river

water
,

1956)

adjacent Yurgenson;
regions

forest

(Sludskii

reservoir

(Shilova-

1940)

1953)

(Kaletskaya, 1957)

(Zharkov

Krasova,

and
Teplov,

1951)

1939)

Mammals
Rodents
Birds

70.6

54.2
53.6
2.3
2.1

57.5

17.1 17.1

34.1

23.6

No

data

69.0
27.0
3.2

56.6

29.8
3.5

20.8
8.3

14.5

Bird eggs
Reptiles

4.0

0.6
8.6

Amphibians
Insects

26.3

8.8
86.7
17.7

2.6

22.5
95.4
36.9
0.1

4.1

17.6
15.3

2.2

12.5

31.9

24.6 88.2

96.0

76.8
18.4
0.8

98.0
9.3

94.4

97.0

Beetle larvae

Molluscs
Plant material

17.7

15.0

23.8

28.3

23.9

221

14.6

144

0.6

12.3

50.7

15.0

77.6

Sample

size

126

474

113

224

134

1953). In the Crimea,

among

insects in the badger's food,

is

ground

beetles and stag-beetles predominate; a large role

fruits

played by

of dewberry, wild pear, apple, acorn, and beechnut.

The

badger also raids vineyards (Savina, 1940). Seasonality of food types is conditioned by the degree of their abundance and availability, which is not the same throughout the
year.

The seasonal changes

Moldavia are

in

food composition of the badger

in

as follows (Korchmar', 1962;

occurrence):

First half of

summer

(572 excrement samples)

Second half of summer-autumn (226 excrement samples)

Mouse-like rodents
Small birds
67.5
7.5

Mouse-like rodents
Subterranean
Stag-beetles
^

18.3

k'oles

10.4

45.2
45.2
31.3

Ground beetle Rose chafer

Beetle larvae

33.3
[P.

Ground beetle Rose chafer Beede larvae Plum

10.0
10.0
7.0

65.0

Sweet cherry
Strawberry
Pear

avium] 60.3
12.9

Dogwood
Apple

44.0
10.0

5.0

Grape

7.0


1268
In Zhiguli (P.

Yurgenson), dung beetles are most frequently

food in May, but in June twice less; later Land molluscs are most often found in JulyAugust, as is hazelnut. Red-backed voles [Clethrionomys rutilus] and wild apples are utilized most of all in August. Mountain leafhopper larvae are more encountered in the badger's food in

found

in the badger's

they rarely occur.

June, beetle larvae in June-August, and wasps (3 species) in July-

August. All of
natural

this

is

associated in a very close

way

with the

It that during one hunt it catches one food type predominantly, but in considerable amounts: thus, in the stomachs of three badgers caught on 3 November, were 57, 63 and 75 frogs (A.N. Skorodumov); in one caught on 31 May 97 maybeetle larvae (Likhachev, 1956), on 27 May 65 maybeetle larvae and 50 dung beetles, on 26 September 285 gm of wasps and their larvae (P.B. Yurgenson), on 26 June 146 barbel beetle larvae, on 5 July 362 larvae of bumblebees, and on 22 August 152 earthworms (Likhachev, 1956). In Moscow district near Klin, during autumn, the stomachs of several adult badgers killed in the

abundance of these foods. is characteristic for the badger

842

morning when they returned to the burrow after a night's foraging, were full of earthworms only, or with small terrestrial frogs only. The latter were, evidently, taken from hibernating aggregations (V.G. Heptner). In Central Europe (Korig)*, 9 old and 74 young voles and 7 frogs were found in one stomach, etc. In general, the badger most often eats not more than 0.5 kg food per day, and young (up to one year old), more than adults. Therefore, an adult badger with an average weight of about 15 kg eats daily a quantity
of food equal in
In the
all

to

3-4%

of

its

live weight.
in

badger are clearly revealed differences

fish,

foods in dif-

ferent biotopes. Thus, in Tataria and adjacent regions, amphibians,

gray voles,

briar,

blackberry and orthopterous insects prebeetles, red-backed voles

dominate

in

foods of animals living in water-meadows; in pine

woods

in broad-leafed forests

maybeetle, click dung

and blueberry;

beetles, mouse-like rodents, earth-

worms and acorns (V.A. Popov). The

Tataria, in

significance of the badger in

is

destruction of the harmful rodents and insects

quite noteable. In

young pine woods, up to 60 thousand places dug by the badger were located, which here destroys up to 75% of all maybeetle
larvae (Gorshkov, 1964).
*Not
in Lit.

Cit. Sci. Ed.

1269

842 Fig. 303. Complex old burrow of European badger in mixed fir-larch forest. South of Vyazniki, Vladimir district. October 1964. Photograph by V.G. Heptner.

Home
away from

range. Almost no data.

its

The badger, apparently, moves

most active

burrow

a considerable distance, but is

within a radius of about 400-500 of about 100 h. Isolated ranges,

m
it

from

it;

i.e.,

within an area
food. With

seems, occur only in those

little

places where badgers are few and the area yields

abundant food, badgers

lies in

live near

each other, sometimes 2-3 fami-

exist in

one burrow of complicated construction. A series of cases which the inhabitants of one burrow visit a neighboring
live together (Neal,

one and peacefully

1948). In Moldavia, in

early spring and late autumn, the radius of badger activity equals

2-3 km;

in

summer and

to 0.5-1.5

km

the beginning of autumn, this range shrinks (Korchmar', 1962). In Tataria, in early spring, the
it

badger feeds near the burrow, in summer, it moves away from for 2-3 km, maximally for 5 (Gorshkov, 1964).

Burrows and

shelters.

The badger

is

a typical

burrowing ani-

mal. Only in very extreme cases, with insufficient places suitable

for burrowing, does
843
it

creep in winter into a haystack or into a hay-

loft in a forest glade. In

England, there was a case where a badger

it

constructed a surface nest-lair in bushes, where

gave birth

to 5

young.

1270

843

Fig. 304. Entrance into

at

the trunk of a

Amur badger, made among stone blocks "Kedrovaya Pad'" preserve, southern Primor'e. June 1962. Photograph by A.G. Pankrat'ev.
permanent burrow of
pine.

Korean

one burrow fluctuates from 1-2 to 4450 (Neal, 1948). In the vast old underground constructions of this animal badger "cities" several badger families sometimes settle. In this event, most frequently, several systems of passages and nesting chambers occur, most often isolated from one another. Usually, some exits are continuously used, the remaining are only

The number of

exits in

used

in case

of danger, or serve for the play of young badgers. In

Tula abatis (Likhechev, 1956), 68.8% of the breeding burrows had

from 2

to 5 exits.

Their width

at the

base

is

from 22

to

63 cm,

height^ 14-32 cm; they are more often semicircular (arched) form. The nesting chamber is positioned 5-10 meters from the exit opening, at a depth of not less than 1 m from the soil surface, more

1271
it is even 2-3 deeper up to meters. General length of the passages varies from 35 to 81 m, with cubic capacity of the entire construction ranging from 8.3 to 19.2 m^. In one case, the volume of the freshly thrown-up ground near the burrow was 38 m^ (Moscow dist.). Average dimensions of the nesting chamber are 74 x 76 cm, with an average height of 38 cm (Likhachev, 1956). Bedding consists of dry leaves and grasses, which are periodically renewed. Leaves are always collected on the dry days. There occur 2-3 nesting chambers. Some of them are open from both ends, and others are blind. In some complicated burrows the system of passages and chambers

often at a depth of about 1.4 m, but in individual cases,

form certain layers ("floors"). Special "ventilation" passages do not occur in badger burrows. In winter, the openings are closed by ground and leaves. By winter, up to 5 kg leaves and grasses for the bedding are collected and the temperature in this bedding is kept constant. The soil at the depth of the nesting chamber most often
does not freeze (Likhachev, 1956).

844

Fig. 305. Occasional day lair of a badger, secluding itself after a night hunt, under a small twisted aspen. Boggy dark aspen wood. In the background of the photo is a small
lake.

South of Vyaznikov, Vladimir

district.

October 1964. Photograph by V.G. Heptner.


1272
Spring cleaning of the burrow is connected with birth of the young (Neal, 1948). Each year, apparently, a new nest chamber is dug for the birth. During the course of the summer it is changed, sometimes several times, apparently in order to rid it of ectoparasites. For the same reason, periodic cleaning of the burrow occurs. The main work of building a burrow dates to AugustSeptember, when a newly formed pair of young dig a new burrow, and old burrows are prepared for hibernation. Daily activity and behavior. As a rule, the badger is active at twilight and at night. On bright moonlit nights, activity drops sharply. Time of emergence from the burrow clearly coincides

throughout the entire year with the beginning of the twilight (a

and changes proportionately with day length Only in extremely remote places does the badger (Neal, 1948). emerge from the burrow during the day, but does not leave it it
little

after sunset),

usually lies in the entrance and basks in the sun.

Among
it

the sense organs, the best developed of

all is

smell

main organ of orientation. The badger's vision is monochromatic. The badger does not react to a lantern with red light. Only moving objects attract the badger's attention. Its hearing is no sharper than that of the human. Since it makes significant noise while moving itself, the badger cannot catch the simultaneous movement of a human. In capturing food, it is mainly oriented by smell. Torpor, winter sleep. Winter sleep in the badger is the adaptation of an omnivorous predator to existence under severe climatic conditions with cold, snowy winter, when its main foods disappear or become unavailable. Winter sleep in the badger, as also in the bear, is not accompanied by a lowering of body
is

the

temperature and slowing

down

of

all vital

functions.
to

Already

in the

second half of summer, the badger begins

accumulate fat reserves, which reaches its limit in October. By this time, the burrow is cleaned and the nesting chambers are filled with bedding. In the middle zone of the USSR, the badger retires
for winter sleep in the period

the middle of the

845

November.
very

It

from the second half of October to usually stops leaving the burrow after
winters, badger tracks are observed

snow
to the
is

falls. In

warm

up

end of January (Tula abatis; Likhachev, 1956). Where

not severe and almost without snow, for example in the
at all

winter

Trans-Caucasus and England, badgers do not retire sleep is repeatedly interrupted, especially by thaws.

or

its

1273

845

temporary shelter of Amur badger Fig. 306. Hollow next to root of Mongolian oak during the period of fattening on acorns. "Kedrovaya Pad'" preserve, southern Primor'e. 20 October, 1964. Photograph by A.G. Pankrat'ev.

Upon

retiring, the

badger blocks the exits of the burrow with

dry leaves and earth.

emerges from the burrow in March-first is 10-15 March, with fluctuation during the multiyear period from 3 to 21-23 March (Likhachev, 1956); in the neighborhood of Gor'ki (A.N. Formozov) the middle of April. The earliest emergence near Kazan was recorded on 2 April (1955) the average date mid- April
In spring, the badger

half of April. In Tula abatis, the average date

1274
has been noted that the time of the badger's emergence from the burrow coincides with the spring arrival of the white wagtail and the beginning of the nuptial flight of woodcock; however, this coincidence is not always observed. In Middle Russia, the badger usually emerges after this, when the average temperature remains for several days above 0C. The short warm spells in the beginning of March do not induce emergence of the badger (Likhachev, 1956). Seasonal migrations and transgressions. Annually, in autumn young badgers disperse. Their dispersal is closely connected with breaking up into pairs and the construction of new burrows. The animals sometimes occupy burrows the inhabitants of which have died. The distance to which young badgers dispense depends upon the food resources of the land. In one case, a pair settled 1.5 km from the parental burrows (Neal, 1948).

(Gorshkov, 1965).

It

Beside such local mixing, cases occur when individual animals

appear
at a

considerable distance from the primary habitat. Thus,

in 1951, a
it

badger was found

in the

upper Pechora, where earlier

are not frequent, but

was unknown (Teplov, 1960). Such cases

are observed everywhere.

How

badgers are able to penetrate for

such distances during their dispersion and what stimulates them to

do

this are not

known.
is,

Reproduction. Reproduction of the badger

time,

at the

present

much

better studied than in the majority of animals having a

still

latent period of pregnancy. Nevertheless, all data are

contra-

dictory.
846

In the

Moscow
it

zoo, young males copulated at the age of one

these matings are productive (Osmolovskaya, 1948). In England, badgers have already united in pairs at the age of one year, but offspring do not occur in the first year, and the first productive mating is observed at the age of 17 months (Neal, 1948). In the G[erman] D[emocratic] R[epublic] and Sweden, adult males are able to fertilize females from January
year, but
is

not

known whether

to October, but yearling males, not earlier than in

March-April or

May

(Shtive, 1952*). Testes of these males are found in active

condition from mid-March to August (other data from

contradictory).

Sweden

are

Females are capable of reproduction at the age of two years (Osmolovskaya, 1948). On attaining sexual maturity in the second
*Not
in Lit.

Cit. Sci. Ed.

1275

"W *"

' ^-

846

Fig. 307.

Temporary burrow of Amur badger

in

oak wood on south slope. "Kedrovaya

Pad'preserve, southern Primor'e. 20 October, 1964. Photograph by A.G. Pankrat'ev.

year,

young females first ovulate either early in spring or in summer and autumn (Neal, 1948). In young strong females, follicles

develop during the period of winter repose (in weaker ones from January to May and later; Unger, 1955). In the middle part of the G[erman] D[emocratic] R[epublic], some of the females born in a
given year are fertilized in September-November, and in the particularly warm winters, also in December-January. In Sweden,

1276
847

the females of that age are not fertilized (Shtive, 1952). Maturation

of follicles in the ovaries of English badgers was observed

February-March, as well as mature, recently ovulated follicLarge, maturing follicles were also noticed in June and September-October (Harrison and Neal, 1956). Ovulation and formation of new corpora lutea also occur in pregnant females during the whole latent period, but esterus,
in
les.

copulation and fertilization do not take place. Therefore, the

number

of corpora lutea

may be

considerably larger than of blastocysts in

the horns of the uterus. During the latent period, not less than

three ovulations take place. Ovulation proceeds within 72 hours

provoked by copulation (Notini, 1948). For implantais no need for the stimulation of copulation. The female allows the male access to her only if it has ready mature follicles. In middle Sweden, as shown by materials from several hundred badgers, females are fertilized from May to July. Copulation in July was also observed in Central Europe, and in general here, cases of summer mating are known to be quite numerous (Unger, 1955). In England (Neal, 1948), copulation was observed in March-May, as well as in June-September, and attempts of it are in February, March and the beginning of October. The greatest sexual excitement occurs in February-March. Rut and copulation proceed for short periods, between which the mutual interest in sex disappears. The least sexual activity occurs in May and up to mid-July, when all badger activity is concentrated on food.
after being

tion of blastocyst, there

In relation to the ability of the lactating females to

lized,

be

ferti-

data are

somewhat ambivalent. According

autumn or
in the

to

some data

(Shtive,

1952)*, they cannot be fertilized during this time, and

fertilization occurs in

following year. The influzoo, estrus and copu-

ence of lactation on the ovulatory cycle has not been shown

(Harrison and Neal, 1956). In the
lation in

Moscow

Amur and
in the

"sand" (steppe) badgers were observed pre-

dominantly

end of March-April, 3-6 days after parturition, and in individual cases in July (Osmolovskaya, 1948). Material of 120 badgers from France showed that, in the south of

the country, in

80%

of the cases copulation takes place during

February, just after parturition (Canivenc, 1957).

*Not

in Lit.

Cit. Sci. Ed.

1277

*^1^'

?'.1^1''

847

Fig. 308.

Track of a badger, emerging early from hibernation, on deep snow. April. "Stol'by" preserve near Krasnoyarsk. Photograph G.D. Dul'keit.

In badgers, there are

two periods of pregnancy:

a prolonged

period of the latent condition of the fertilized egg, and a

shorter period of active development of the
tation of the

much

embryos

after

implan-

ovum.

In the latent period, the fertilized eggs reach

the blastocyst stage and are found free in the horns of the uterus.

Their nutrition

at this

time

is

probably fulfilled by the secretory

activity of uterine epithelial cells. In the period

March-July, the
is

blastocyst has a diameter from

primitive trophoblast.

1.5
to

to

2.0

mm this

the

From October

December, the blastocyst

already has a diameter of 3.5-4.0

848

mm.

Their placentation proceeds

December, when they are evenly distributed along the uterine walls (Canivenc, 1957). In England (Harrison and Neal, 1956), blastocysts were found from 30 April; from placentation to parturition takes 60 days. In general, the duration of pregnancy the case of summer mating is 271-284 days, and for early spring up to one year. In
in

1278
females fertilized in January and
the end of
later, the latent

period extends to

December, and the duration of pregnancy consists of 14-15 months (Shtive, 1952)*. With castration and ovary ligation, it was shown that up to

implantation, hormonal secretions of the ovary, including the

at ail on nutrition and development of the blastocysts (Unger, 1957). Introduction of proestrone and progesterone with estradiole did not stimulate

corpus luteum hormones, have no effect

placentation.

It is

possible that the latent stage

is

the result of the

absence of the element necessary for

this process,

perhaps ovarian

hormone. The quick development of the embryos after placentation is coincident with the lengthening of day light; i.e. with the effect of cortical hormones (Unger, 1955).

The average number of corpora

was

lutea in material

from England
in Janu-

(Harrison and Neal, 1956) during the entire period of pregnancy

6.5 (from 2 to 11), the average

number of embryos

ary-February

Germany 2.7 (Fisher, 1931).* In England, the average number of young in a litter is 2.25 (1-5); in 22 cases, it was 2; in 8 cases, it was I and 3, and in 4 cases 5 (Neal, 1948). In Moscow zoo, the number of newborns ranged from 2 to 5 (average 4). In Kazakhstan, in the Dzhungarsk Alatau, litters were 2-4, in the lower Hi 3-4, and in Tselinograd district 5-6 (Sludskii, 1953). On the whole, the size of litters varies from 1 to 6 and the number of embryos reaches 7, but most often in the litter 2 badger

3.1; in

Sweden,

it is

equal to 2.5 (Notini, 1948); in

whelps occur.
In the G[erman] D[emocratic] R[epublic], young are born in March-April; in Sweden in March; in Western Europe, from

19

December

to

14 April. There was one record in July (Neal,

1948). In southern England, [birth] occurs

more often

in in

February-

March and
the

in the north of the country

more often

March. In

Moscow

zoo, in 7 cases parturition occurred in the period

26 March-21 April. Therefore, the reproductive process in badgers quite complicated. The major periods depend on geographic variations in living conditions. These also have, undoubtedly, significance also for age, and attributes of early or late litter, etc. In nature, badgers are monogamous, and most often the female copulates with one male; however, rare cases of mating with two
is

*Not

in Lit.

Cit. Sci. Ed.

1279

males have also been observed. The male remains with the female
for several years, possibly for
its

entire life (Neal, 1948).

Growth, development and molt. In England, the newborn badger pups weigh about 85 gm, with body length of 12 cm (Neal, 1948). Weight of newborns in the Moscow zoo was 63-84 gm; on averears covered

age 75 gm. They are born completely helpless, blind, and with the by skin. The newborn cubs are covered with sparse,
white wool and the dark stripes on the head are well differentiated
indiat

At the age of 3-5 days, the claws become pigmented and

vidual dark hairs appear. Strong darkening of color
the age of
is

observed

849

10-14 days. The juvenile pelage is replaced at the age of 41-68 days. Ear pinnae are formed at the age of 3 weeks, and eyes open on the 35th-42nd day. Milk teeth erupt at the age of 1 month, but are not fully developed, and at the age of 2.5 months, their replacement by the permanent ones begins. Reduction of the milk dentition may be the result of prolonged feeding exclusively on maternal milk. The young shift to independent feeding at the age of 3 months (Osmolovskaya, 1948). At the age of 4-6 months, a reduction in intensive growth is observed. At the age of 6-9 weeks, badger pups begin to emerge from the burrow, but they do not decide to abandon it. The first emergence under the mother's protection was noted in England at the
end of April. At the age of 11 months, they come out near the burrow to play, but only in the presence of parents. Dispersal of young badgers proceeds in October, and at that time, they break up into pairs. Enemies, diseases, parasites, mortality, competitors and population dynamics. The badger has almost no enemies. Wolves, lynx, and dogs are dangerous to it, but cases of death from them are very
rare.

In practice, only

human

activity has significance for the

it from occupied areas by destruction of burrows. This latter is particularly severe where possibilities of burrowing are very restricted because of unfavorable ground and hydrological conditions.

badger

direct destruction,

and also forcing

In nature, cases are

known of

a die-off

among

badgers, after

which the animals disappeared from the given

the die-off were not established.

locality for a series

of years. This points to the effect of an epizootic, but the cause of

spreading in badgers

pulmonary

Only an invasion of helminths known. The most dangerous of these are invasions, in particular eluerostrongylosis and
is

1280

h m %

toA

(It

849

Track of right paw of badger on moist sand and the scheme of tracks during slow movement of the animal. Naurzum preserve, Kazakhstan. 7 April 1957. Sketch by A.N. Formozov, about 2/3 natural size.
Fig. 309.

krenozomatosis (Rukhlyadev, 1940). In England, mortality due to

acute tonsillitis was recorded (Neal, 1948).

The extent of mortality

is

among badgers

is

unknown. The age

limit

supposed to be deter-

to burrows the badger's competitors are shown to be the red fox and raccoon dog. However, the badger is by no means appears here as the weakest. The old, widespread represen-

mined as 12-15 With respect

years.

tation that the fox itself, being not purely carnivorous, drives out

badgers from burrows,

is

untrue. Often, fox and badger live side

by side
its

in isolated sections of a

cases are
litter,

wide burrow. However, a series of away a fox and destroyed discarding the fox carcasses on the surface (Likhachev,

known where

a badger drove

1956).
In England, badgers often die under the wheels of cars

and

trains at night, but in the majority of cases, they apparently die in

burrows. This

is

supported by the frequent discovery of skulls and

bones

in

seal the

burrow scrapings during cleaning. Apparently, badgers chamber in which the badger died and dig a new one in

another part of the burrow. In England, they observed, as a pair of


1281

badgers pulled a badger corpse from a burrow and buried

rabbit

it

in a

850

burrow widened for this (Neal, 1948). There are fluctuations in badger populations, but have not yet been studied. The greatest influences on them are rendered by humans and diseases. In England, for example, a considerable increase in their number was recorded in the war years in connection with the cessation of its pursuit by humans (Neal, 1948). Field characteristics. In twilight and night time, the badger can be detected in the forest by its noise made while moving and feeding. Under the cover of darkness in distinction from the majority of wild animals, it does not conceal itself, and is heard from a far. Characteristic signs indicating the presence of the badger are the beaten tracks going in various directions from its burrow to a distance of up to 0.5 km, numerous, very typical holes made by it while catching mouse-like rodents, beetles and their larvae, destruction of bumble bees and wasps nests, and also by overturned forest litter, and feces left here and there. Tracks of badger on dirt, moist soil or on sand are difficult to confuse with anything else. They resemble most of all tracks of bear in miniature. Prints of the plantigrade feet and long, massive
claws are particularly characteristic of them. Most often, prints of prints of hind and fore paws overlap the tracks occur in pairs

each other. Length of the forefoot

and of hind about 9 cm; length of claws of fore paw in prints is about 2.5 cm, length of doubled prints of the track, about 10-14 cm (Formozov,
is

about 8

cm

1957).

Practical Significance

economy, the badger has little, mostly local, significance. The price of fur product from it is not high. The skin is used for shagreen [leather] and the guard hairs for making shaving brushes of high quality. Badger meat is eaten in several districts; however, in the majority of cases, it is thrown away. Badger fat is highly valued, in Middle Russian districts (Moscow, Vladimir) for example, and the badger is mainly hunted for its fat rather than its less valuable skin. The fat is used for greasing leather, but it is
In the hunting

chiefly used in folk medicine in case of injuries, rheumatic dis-

eases and even tuberculosis;

i.e.,

analogous to bear

fat.

On

the

market,

it

is

very scarce (V.G. Heptner).


1282

Not less than 50% of the yearly catch of skins is obtained from Kazakhstan and other Middle Asian republics of the Union. Here the badger constitutes about 5% of the commercial fur trade and in other places considerably less. At the same time, the badger is useful in destroying insects
and mouse-like rodents harmful to forestry and agriculture. Among insects and their larvae that are destroyed in great numbers are
larvae and adults of maybeetles, larvae of mountain leafhoppers,

mole

crickets, locusts, larvae of

longhorn beetles and sawflies.

vole has greatest signifi-

Among
cance in

mouse-like rodents, the

its

common

food, and pests of forestry and agriculture.

The badger
very insig-

feeds also on injured animals and birds, and on their carcasses.

Harm caused by
nificant.

the badger to the hunting

economy

is

The accusation of badger

attacks on small domestic live-

stock

is

not confirmed. In England such cases in relation to domestic

fowl were observed, but very rarely.

Methods of hunting the badger

vary, except that the badger

does not serve as the object of special hunting. Badgers are caught in jaw traps, or opportunistically by night hunting with dogs, am-

bushed

burrows with guns, and also caught with special dogs and dachshunds) used to excavate the burrow. Finally, the badgers are sometimes smoked out of their burrows, but more
at

(fox-terriers

often, they die in the burrow.

Calculated against the useful activities of the badger,

in the

its

taking

zone of broad-leaf forests and forest-steppe and in the regions of field-protecting forest belts should be entirely prohibited throughout the whole year, as also should, any kind of excavation and damaging its burrows, especially where the number of badgers is determined by the conditions for burrowing. Spring and summer hunting should be prohibited everywhere as well, except in the

autumn months: from 15 August or

September

to

November (P.Yu.).


1283
851

Subfamily of Otters
Subfamilia Lutrinae Baird, 1857

Genus of Otters

Genus Lutra Brisson, 1762

1762. Lutra. Brisson. Regn. Anim., 13. Mustela lutra Linnaeus.

Dimensions

large.
light,

General construction

body strongly elongated and

tail

thin,

extremely flexible; head small, small ear pinnae;

long, exfin-

tremities strongly shortened, plantigrade, claws small,

between

gers webbing strongly developed. Pelage very close-fitting, with

coarse guard hairs and very dense, soft underfur. Color almost
constant over whole body, more or less lightening on lower surface, of

tawny tone. Glandular area

at

base of

tail

absent. Teats 2

or 3 pairs.

Skull relatively light, broad and strongly flattened, with strongly

diverging zygomatic arches and very sharply narrowed postorbital

region. Braincase
its

volume enlarged,

rostral region short

and broad

width greater than length. Posterior part of hard palate lying

posterior to toothrow narrow and relatively short. Mastoid proc-

esses flattened, approaching anteroventral part of occipital crest,

paroccipital processes

low but separated from auditory

little less

bulla.

Mastoid width of skull a

flattened, with small

than zygomatic. Auditory bullae

bony auditory meatus.

Dental formula
dentition
is

C-P M = 36.

14

In

its

main

features, the
It

like that of the

badger subfamily, Melinae.

also

represents the

same combination of
to the general

cutting and tubercular (grind-

ing, crushing) type, with

however, great development of incisor

dimensions of the
is large.

characters.

Compared

rostral part

of the skull and of the palate, the dentition

sial

Upper carnasin

tooth large and triangular in form.

its

Upper molar rhomboidal

form,
less.

area approximately equal to carnassial or very slightly

carnassial tooth long, anterior part has three welllittle

Lower

developed apices; posterior part a

longer than anterior. Loss

of anterior premolars usually does not occur.

1284
Specialized freshwater forms, excellent

swimmers and

divers,

always connected biotypically with water bodies, both in plains and in part, mountains. Swims both by paddling movement of the
limbs and by sinuous bending of extremely flexible trunk and
tail.

Feeds mainly on

fish, in part

on aquatic invertebrate (crayfish) and

is

amphibians. Builds bank burrows. Hibernation

ently, as is a latent stage in

absent, appar-

development of the
is
It is

fertilized egg.

The range of
very largest
several bats. This

the genus

extremely extensive

one

of the

among mammals.
is

exceeded only by the ranges of

particularly remarkable, given the great spe-

cialization of species in the

genus and the relatively small number

is

of them (see below). The genus

distributed throughout Europe,

occupies the mainland of Asia (except the extreme north and

Arabian Peninsula), Ceylon, Sakhalin, the Japanese islands, Taiwan, Hainan, Java, Sumatra, Calimantan (Borneo) and Banka. Representatives of the genus are absent on the remaining islands of the
Pacific Ocean'.
853

In Africa, the range includes the extreme northwest of the

continent (Atlas countries) and the entire continent south of the

Sahara, except its extreme southern and southwestern parts. In North America, the range occupies the whole continent, except several areas in the extreme north and desert regions in the south,
the islands adjacent to the continent along
its

western shore (Kodiak,

Alexander archipelago; absent on islands of the Bering

islands off the northeastern shore
ones). In the range
is

sea), and (Newfoundland and some smaller included Central America and South America,

and the whole of Tierra del Fuego. In the range is included the island of Trinidad, but species of this genus are absent on all
islands of the

West

Indies.

The range of

the genus, therefore,

occupies almost the entire range of the subfamily

absent only on Palawan,
lands, in middle India and in South Africa,

its

species are

Commander, Aleutian and

Pribiloff is-

where species of other

it

genera are encountered.

The genus Lutra

to

is

the very largest in the subfamily. In

are

11 species of the 16 assigned to Lutrinae.

divide

it

There were attempts into several independent genera (Lutrogale for

incorrect (see below, in description of

851
otter).

'Information on the Kuril islands

is

Pocock's reference to its occurrence in the Philippine Islands is, apparently, wrong it was not confirmed by anyone. In the Philippine sensu stricto, true otters are absent, but the clawless otter, Aonyx cinerea, exists on Palawan.


1286
perspicillata,

L.

or Hydrictis for L.

maculicollis)\ however, the

it

majority of authors do not accept this division, and

ently,

is,

appar-

more correct

to

recognize one genus with two subgenera

Lutra and Lutrogale.

On

the other hand,

some authors (Chasen,

1940; Carter, Hill and Tate, 1946) believe that the separation of
the genera (subgenera) of the so-called clawless otters

Aonyx
basis.

(capensis) and

Amblonyx (= Microaonyx: cinered) has no

They combine the species mentioned with Lutra, which is an hypothesis that demands a wider foundation. Besides the most widely distributed Palearctic and IndoMalayan species, the common otter, L. lutra, the genus also includes the following species: L.* canadensis (North America, south-

ward

to

Mexico)^;

L.

sumatrana (Indo-China and Malayan

archipelago; evidently, the least specialized species); L. maculicollis

(Africa south of the Sahara); L.* annectens (extreme south of North

[America], Central and South America); L.* enudris (South America); L.* felina (southern South America; a marine form, evidently the most specialized form in the genus); L.* incarum (South America); L. * platensis and L. * provocax (the same;
L. perspicillata (South and Southeast Asia; subgenus Lutrogale). Therefore, the greatest richness and diversity of species are obviously, characteristic of South America (the center

subgenus Lutra) and

of origin)

there
1

are 6 species out of

1 1

here. North

America
1

is

characterized by

species, there are 2 species in South Asia (one

1

represents a separate subgenus),

species in Africa and

species

has the above-mentioned wide distribution in Eurasia and northIt is noteworthy that with the great diversity of South America (besides those mentioned, there is also the giant river otter, Pteronura brasiliensis). North America possesses only one endemic species. One is characteristic also for northern Eurasia apparently the diversity of forms in the genus is in the

western Africa.

otters in

south. In this sense, geographic variation of L. lutra

tive (see below).

is

also indica-

As already mentioned (see characters of the family), in this complicated mix of forms representing the marten family,
*These

New World

species, here placed in Lutra, are

now

considered a distinct

genus, Lontra

Sci.

Ed.
its

^This form, with

a

great general similarity to the Eurasian otter, itself represents

characterized craniologically. The separate species, mitra (Prince Wales Island in Alexander archipelago), accepted by American authors, represents only a subspecies of L. canadensis.
separate
species,

well

L.

1287

Mustelidae (perhaps
the otters

itself a
is

polyphyletic group), the subfamily of

Lutrinae

Perhaps

it

may even be

one of the most accurately circumscribed. considered the most natural and sharply

distinguished one.

Besides the genus Latra, the subfamily includes 4 other genera, the so-called giant river otters:
(1

Pteronura, in South America

854

Aonyx a genus two species belonging two subgenera comprising to A. (Aonyx) capensis in Africa and A. (Amblonyx) cinerea in Southeast Asia, Paraonyx with 1 species in West and Central Africa {P. congica: "species" microdon and philippsi are not taken into consideration Enhydra, with 1 species {E. lutris) here) and sea otter* (kalan) from the shores of the northern part of the Pacific Ocean (see
species
P. brasiliensis), the clawless otters,

below).

The
genera

identification of the position of Lutra within this circle of

is

very difficult. The thing

is

that within the limits of the

subfamily, the genus of sea otters ("kalan")

sharply set apart from
all

Enhydra

is

totally

the genera of the subfamily by the exits

tremely unique structure of

teeth

limbs, skull, completely distinctive

and other characters (see description of this form below). All the remaining genera (and species), together and separately, are very far different from Enhydra and are, to a greater or lesser
degree, close to each other. Differences between them are completely unequal to differences

between each of them and Enhydra.

By

this criterion,

they might perhaps be considered as one genus;

so to speak, river otters, Lutra; corresponding to the genus of sea

otters,

Enhydra.

Characters of external structure and craniology, by which the

otters are usually grouped, create

patchwork combinations. Thus,

capensis) or they are very small

the group

Aonyx has no claws

is

(A.

(.

cinerea). This [latter]

usually considered a specialized charis

acter ("progressive").

At the same time, finger webbing

reduced

or actually almost absent as in the posterior limbs of A. capensis.

Their skulls are relatively short with a voluminous braincase. The

dentition
is

modified only weakly and unsubstantially. As regards

its

the structure of

skull,

extremely developed, large finger web-

bing and certain other characteristics, evidently the entire genus

Lutra must be considered a highly specialized ("progressive") group

*In Russian original,

"Kamchatka beaver"

Sci. Ed.


1288

among

the genera of freshwater otters.

is

Most

closely related to

it,

apparently,

the genus of giant Brazilian otters Pteronura.

The

southern Asiatic and African Aonyx and Paraonyx are, apparently,

more
In

primitive, although
its

Aonyx

is

also very close to Lutra.

origin, the subfamily is apparently

connected to certain

extent with the subfamily of badgers, Melinae. However, the earliest representatives of the subfamily as highly specialized forms

already known from the Upper Oligocene of Europe (Potamotherium). More than 10 genera of fossil otters have been described, mainly from the Upper Tertiary period (Miocene,
are

Pliocene

Paralutra, Enhydriodon, Vishnuonyx, Sivaonyx and partly from the Pleistocene (Cyrnaonyx). The genus and others) Lutra is known from the Lower Pliocene of Europe and Asia and from the Upper Pliocene of America. All the species of the family, even in the tropics, represent
valuable fur animals.
In the

USSR,

there

is

1758, which represents about

one species: Lutra (Lutra) lutra Linnaeus, 0.3% of the species of the fauna and
the

about

10%

of the species in the genus.

The range occupies almost

The species
is

whole

territory of the country.

everywhere closely associated with water.

valuable fur-bearing animal (V.H.).

Subgenus of the

Common

Otters

Subgenus Lutra Brisson, 1762

[EURASIAN] OTTER, PORESHNYA
Lutra (Lutra) Linnaeus, 1758
1758. Mustela lutra. Linnaeus. Systema Naturae, ed. X,
1,

p. 45.

Uppsala, Sweden.
1777. Lutra vulgaris. Erxleben. Regn. Anim.,
1, p.

448. Renaming

of lutra. 1834. Lutra nudipes. Melchior. Den Danske Stats og Norges Pattedyr, p. 50. Coast of northern Norway.
855

1839. Lutra aurobrunnea. Hodgson.

J.

As. Soc. Bengal,

8, p.

320.

Nepal.

1289
1839. Lutra monticola. Hodgson. Journ. As. Soc. Bengal,
320. Nepal.

8,

p.

1844. Lutra kutab. Schinz. Syn.

Mamm.,

p.

354. Kashmir.

1867. Lutronectes whiteleyi. Gray. Proc. Zool. Soc. London, p. 181. Japan. 1912. Lutra lutra seistanica Birula. Ezhegodn. Zool. mazeya Ak.

nauk, 17, nauk, 19,

p.

274. Gil'mend river in Seistan, eastern Iran.

muzeya Ak. XXI. Lyangar, Pyandzh basin. Western Pamir. 1922. Lutra vulgaris var. baicalensis. Dubowski. Arch. Tow. Nauk. Lwow, 1, p. 349. Nom. nud. Near Baikal. 1922. Lutra vulgaris var. amurensis Dybowski. Ibidem, p. 349.
1915. Lutra lutra oxiana. Birula. Ezhegodn. Zool.
p.

Nom. Nom.

nud.

Amur and

Ussuri Territories.
p.

1922. Lutra vulgaris var. kamtschatica. Dybowski. Ibidem,

nud. Kamchatka.
1931. Lutra lutra meridionalis. Ognev. Zveri Vost. Evropy
Azii, 2, p. 527. Vic. of Teheran, Iran.

349.

Sev.

1936. Lutra stejnegeri. Goldman. Journ.

Mammal.,

17, p. 164. Vic.

of Petropavlovsk, Kamchatka (V.H.).

Diagnosis

The only species of

the genus in the

USSR.

Description

Body

strongly elongated and relatively thin,

its

more or

less equally

thick along

whole length

posterior part only slightly

more

massive than shoulder region. Limbs short, from whence its stature. Tail long and comprises about half the length of head and
body, very thick
at base, strongly
tail

narrowing to
structure
all
is

tip

and flattened

in

dorso-ventral direction. Such

determined by strong

development of ligaments but most of

cular
tail tracts

by the powerfully mus-

running along the vertebrae. Moreover, masses of

less

collagenous tissue are strongly developed along the lateral sides of

the
tail,

somewhat

on the lower

side,

and

still

less

on the

upper
fore,

side, forming loci for fat deposition. Tail thickness, theredepends to a certain degree on nutrition of the animal, and
it.

characterizes

The

tail,

contrary to

all

remaining species of the


1290
family (except sea
otter,

does not bear long hairs.

It is

covered by

short (as on the trunk or shorter) closely investing fur.

Head

relatively small, elongated oval, narrow,

no wider or
fur,

hardly wider than the neck, strongly flattened.

Neck

relatively short.

Ears short, with rounded apexes, weakly protruding from

widely

separated. External ear openings covered by special valves. Eyes

quite large; bare area on tip of nose large, transversely elongated
its

width greater than

its

height or approximately equals to

is

it.

On

the whole, the appearance of the otter's head

somewhat

snake-like.

Digits fused together by strongly developed webs, reaching

middle or nearly

to

end of

distal phalanx. Digits, including first,

somewhat elongated, which increases area of hand and

foot.

Second and third digits on both fore and hind limbs are longest, and equal in length. Lower hand and foot naked, edged with elastic hairs somewhat increasing propulsion surface. Claws small, but
relatively
little

reduced.
is

The
active.

otter

body

extremely flexible and the animal

is

very

On

land, the otter

moves

in a
tail

walk or

trot,

with lowered
its

head, bent back, and outstretched

856 the

usually dragging
[its

end on

ground; more rarely leaps, strongly hunching

its

back].

On

the

Movements of otter in water are particularly confident and agile. Swimming slowly, it usually paddles with its paws but when moving quickly, it presses limbs against body and moves with energetic snake-like movements of the whole trunk and muscular tail. The moveability and
whole,
is

movement on land

not very agile.

flexibility of the

vertebrae and entire body of the animal are

is

exceptional. Overall, in general appearance and habits, the otter

entirely unique

and

is

strongly differentiated from

all

remaining

species of the families of our fauna.

even over the whole body (difference between length of hair on back and belly is small), closePelage
in

winter

is

short, very

fitting, shiny,

with often coarse, elastic guard and very dense and

close-fitting soft, silky underfur.

The

fur on the belly

is

denser

than on the back. In northern otters, the average length of contour

hairs

on the back averages 24.2

hairs, correspondingly, are 18.4

and 11.2

mm.

mm, on belly 21.0 mm; guard and 17.2 mm, and underfur 14.6 The number of hair on 1 cm^ of dorsum reaches

35,000 (34972), on venter about 50,000 (50668). On the back, for every outer hair, an average of 155 underhairs occur and on the


1291

856

Fig. 311. Otter, Lutra lutra L. Sketch

by A.N. Komarov.

belly

120. Outer

hairs

have very wide and flattened

is

distal thirds

the ratio of their width to length

1:3-3.5 (in terrestrial forms,

not

more than

1:1.5-2.5). Moreover, the outer hairs are strongly

inclined with respect to the skin surface (30-35) and are strongly bent near the base ("in the neck") where they are relatively thin.

On

the whole, the outer hair has the

form of

a small plate

and

together they tightly cover over the underhairs.

latter, as

The bases of

these

well as bases of the smallest top hairs, are strongly crimped

coat. All these characters, especially the struc-

and form a wavy fur

an aquatic

ture of the outer hairs, themselves represent typical adaptations to

mode

of

life,

and they prevent wetting of the

fur,

even

during extended immersion in water. The cortical layer of the hair

1292
is

greatly

increased,

thus

its

durability

greatly

augmented

(Tserevitimov, 1958)^.
857
life;

Full growth of the otter

is

attained only in the third year of

at that time, its fur is also

completely developed. Skins of

animals born

in the current

year ("koshlak") have a long and soft

fur

pelage composed of sparse underfur and thin, soft guard hairs. In

second-summer animals, the

Glandular areas

is

longer and softer than in adults,

but coarser than those born in the current year, with sparse underfur.
at the tail

base are absent; anal glands are

present, but not strongly developed (?). Teats

2-3

pairs'*.

Color of the upper body

color as back.

in winter fur is bright

dark-brown or
of the same

dark nut-brown. Upper head somewhat darker,

tail

Lower

surface of body, including ventral neck and

lighter,

lower and

lateral

head are slightly

with tinge of silvery

yellowish tones, sometimes whitish.

a gradual transition
belly.

On

the sides, the color forms

from

the tone of the back to the color of the

light horn, darker at the base.

Legs are dark brown, and claws

In

summer

coat, the fur

is

somewhat more
is

sparse, but of the

in

same color

as in winter.

There are no differences among sexes

color. Individual variability of fur

insignificant and lies only in

more

or less intensity, and in greater or lesser lightening of venter

relative to

dorsum. In some
lighter

localities,

two color types are

differ-

and darker (G.F. Bromlei; southern Ussuri Territory); however, color dimorphism does not occur.^ This is normal, simple individual deviation, apparently, even mostly age variation.
entiated

The

soft "fluffy" fur of the

first

young

otter

is

dark-chocolate
darker

color and lacks luster. At

winter, the

young animals don a

is

coat similar in color to that of the adult, but their venter

than in adults.
to

Upon

attaining the adult condition, color continues

later color

change somewhat, and

changes

in the direction of

'Otter fur

is

considered very durable and "long-wearing" and in determining the

its

relative quality of other furs,

"long-wearing"

is

accepted as 100. In mink,

it

equals

70, in sable 50 and in squirrel 25 etc.

For otters of England, 3 pairs recorded (inguinal; Miller, 1912), for middle European 2-3 pairs (Gaffrey, 1961), for our otters, 2 pairs (Ognev, 1931), for the Indian form of the species, 2 pairs (Pocock, 1941). 'It is considered that the dark otters, called "river", have a uniform dark-brown color and small dimensions, and live in rivers. The other form "sea" otter is larger, its color is lighter with a dirty-ocherous belly and legs, lives and feeds in the sea (G.F. Bromlei). Similarly, two types of color were also noted for otters in the upper

Amu-Darya

basin (Chernyshev, 1958).

1293
lightening of the general fur tone, while the venter
relative to the

becomes

lighter

dorsum. Geographic variations of color within the

USSR

are insignifi-

cant and are only expressed in lightening of the general color tone,
relatively greater lightening of venter,
in

and decrease

in the fur density

some southern regions. The skull is very broad, both

its

in rostral,
is

and particularly

in the

zygomatic and cranial regions, and

the braincase in

strongly flattened. Height of

middle section constitutes only slightly more

than half the mastoid width. Height in the region of the auditory
bullae
file is
is less

than mastoid width.

The upper

line of the skull pro-

almost straight

it is

only insignificantly depressed in front

of the middle of the interorbital area; and in the young

occipital region.
line of the

also

in the

On

the whole, this line

skull.

is

almost parallel to the

skull
is

lower surface of the

Such a flattened

absent in any other carnivore of our country.

posterior parts.

The braincase is very voluminous and broad in its middle and The width of this region is increased still more by
which

strong development of the occipital crest which extends also onto

the lateral surface of the skull, and the mastoid processes, are strongly flattened and fused with

them

in its anterior part.

The
is

anterior region of the braincase (in the interzygomatic area)

sharply narrowed. The postorbital area is sharply narrower and forms a strong intersection, particularly striking to the eye thanks
859 to the

wide separation of the zygomatic arches. Width of the

is

postorbital constriction
interorbital area

considerably less than the width of the

and

is

approximately 4 times less than mastoid

width. Postorbital processes are well developed, but short and blunt,

and border only a very small part of the orbit. Orbit quite large its greatest diameter almost equal to interorbital width. Zygomatic arches are strongly separated laterally throughout their whole extent (as well as in anterior part) and generally form a rounded figure. They are strong and massive, noticeably curved upwards, and in the anterior part bear a well

defined tubercle on their upper edge, bordering the orbit from

behind and below. Infraorbital foramena, perforating base of zygomatic arch anteriorly, very large. Its diameter is 1.5-2 times

more than diameter of

cranial), high

the cusp of the upper canine.

Rostral part of skull very short (about 2 times shorter than

and broad. Distance from anterior point of zygomatic

1294

858

Fig. 312. Skull of otter, Lutra lutra L.

arch to anterior edge of alveoli of incisors approximately equal to

width of muzzle below canines, and approximately 4.5 times less

than condylobasal length of the skull. Sagittal crest
quite narrow and elongated
is

almost un-

defined or only weakly so. Palate between toothrows relatively

distance

between inner parts of

1295

859 Fig. 313. Skull of a very old otter, Lutra lutra L. Posterior cheek teeth only remain, worn to gums. Sudzukhinsk preserve, southern Primorye. Sketch by G.F. Bromlei.

molars

fits

into length of palate

from anterior edges of alveoli of

is

incisors to posterior edge of molars about three times. Portion of

palate lying behind the toothrow

wide

anteriorly, quickly nar-

rows

posteriorly,

and

is

short. Its length is

approximately equal to

distance between canines. Interpterygoid notch narrower anteriorly

than posteriorly.

Auditory bullae small, flattened and only insignificantly swol-

form of short rounded outgrowths and do not contact auditory bullae. The external auditory meatus is short, opens below mastoid (anterior) part of occipital crest, and are directed forward and upward. The lower jaw massive compared to general lightness of skull. Coronoid part short, but quite wide, its apex not pointed. Compared to general dimensions and width of palate and maximum transverse diameter of upper rostrum, teeth are large carnassial tooth constitutes half of distance between carnassial teeth.
len along inner part. Paroccipital processes have

Canines are thin, long and very sharp. All small premolars are well developed and of tearing form, and by their location in relation to
teeth of opposite
tus.

jaw (upper-lower), form a strong tearing appara-

upper premolar well-developed and not reduced. It is displaced deep into the toothrow and does not lie directly behind
First

the canine, but

on

its

intero-posterior side.

Upper

carnassial tooth large, quite regularly triangular in form,

860

one angle directed inward. It bears a high cutting apex along outer edge, behind which lies a second apex, shorter and connected with first by a crest. In front of the main apex lie small teeth. All of them are arranged regularly next to one another. The inner blade.

1296

which constitutes about half the area of the tooth, is and does not bear apexes. Upper molar large, rhomboidal in form. Its area is approximately equal to area of carnassial tooth or slight smaller, and its
the area of
flat

greatest diameter

is

directed almost at a right angle towards the

axis of the toothrow.

tubercles, of

The

flat

surface of the tooth bears four small

which two are along the outer edge. Lower carnassial

tooth elongated in form, with three well-developed sharp apices,

almost identical in dimensions on the anterior part of the tooth.

Posterior half of tooth

somewhat

larger than anterior

one and does

not bear sharp apices. In area, second lower molar significantly

smaller

approximately three times than

is

carnassial.

Sexual dimorphism of the skull

well manifested. Beside someis

what smaller general

als,

size,

the female skull

distinguished by

almost complete absence of a sagittal crest, even in old individu-

somewhat smaller

infraorbital foramen,

narrower

rostral part

of skull (breadth above canines), somewhat smaller nasal opening,

and, apparently,
notch. Skulls of
crests, etc.,

somewhat narrower and shorter interpterygoid young animals have less-developed protuberances,

and are less angular; nasal region is relatively low, lower than occipital region (with age, massiveness and height of
anterior part of skull increases), postorbital region
is

broader and the

supraorbital processes are small (Ognev, 1931). Sagittal crest does not

develop and occipital

within the

is

very small. Geographic variation of the skull

if

USSR

was not noted, and

is

expressed, then very weakly.

el-

Posterior section of os penis

evated.

thickened and somewhat

tips directed

On

anterior end has

two rounded lobes,

down-

ward separated by groove. Left lobe 1.5 times thicker than right and ventrally bears incision (Ognev, 1931). Ratio of body length to length of intestine (two males) 1:6.36 and 1:6.10; cardiac index (male, January) 13.8 (from data of Chernyshev, 1958; Pyandzh basin).

Body

length

foot length

is 550-950 mm, length of tail 260-550 mm, hind 110-200 mm, ear length 17-25 mm.

Condylobasal length of male skull is 107.0-126.0 mm, of females 96.5-121.0 mm; zygomatic width of males is 62.0-83.0 mm, of females 60.5-72.0 mm; interorbital width of males is 17.5-22.5

mm,
18.3

of females 16.2-21.3

mm;

postorbital width of males

is

12.3is

mm,

of females 11.8-18.3

54.5-71.3

mm,

of females

mm; mastoid 55.9-67 mm.

width of males

1297

Weight of
Females, as

adults 5.7-10 kg.^

is

evident from the craniological indices presented,

are noticeably smaller than males. Differences in

body dimensions
(Sudzukhinsk

and weight of individuals from one restricted

mens): body length of males; 550-810

territory

preserve in southern Ussuri Territory) are as follows (15 speci-

mm, of females 550-680 mm, of females 280-380 mm; hind foot length of males, 110-120 mm, of females 82-93 mm; ear length of males, 19-25 mm, of females 17-22 mm. Maximum

mm;

tail

length of males; 280-455

861

9470 gm, of females, 6543 gm. Animal in the first year of life, in November-February, have a body length of about 350 mm and weight from 1100-1500 gm (G.F. Bromlei). The latter figures are relatively small young otters from Pechora (preserve; Z[oological] M[useum of] M[oscow] U[niversity]) in November-February weighed 2250, 2400 and 4430 gm (males) and 2250 and 3500 gm (females). Very old individuals approaching the age-limit experience poorer nutrition than those of intermediate age and, despite sometimes large dimensions, weigh less (G.F. Bromlei). Nutrition and weight change seasonally. Length of OS penis is 58-65 mm. Geographic variation of dimensions within the boundaries of the USSR was not noted, but some, apparently, exists, and southern otters are somewhat smaller than northern. Several forms from
weight of males
is

the extreme south of Asia (nair, barang) are quite strongly differ-

entiated

by

their

dimensions (V.H.).

Geographic Distribution
In Europe, Asia except the

extreme north and Arabian Peninsula,

and in northwestern Africa.

*Data on body dimensions and weight from Ognev, 1931; Novikov, 1956;
Stroganov, 1962; G.F. Bromlei and from materials of the Z[oological] M[useum of] M[oscowj U[niversity]. Skull dimensions from Ognev, 1931; Stroganov, 1962 and
collections of Z[oological]

M[useum

of]

M[oscow]

U[niversity]. Altogether approxi-

mately 150 skulls. Reference to body length of 100

cm and
is

weight of 15 (16) kg

(Central Europe; Gaffrey, 1961 and several other sources)

apparently an exaggera-

tion. In any event, such measurements and weight were not recorded in our otters, even the very largest populations. However, one case is known of an otter caught in Turkhan Territory with tail length (measured from skin) of 175 cm (Zalesskii, 1930). It is possible that this is a case of gigantism, analogous to kolonok and polecat (see

[above]).

1298

Geographic Range

in the Soviet

Union
part of the range of the spe-

Very vast and constitutes the greater

cies. It

occupies almost the entire territory of the state except the

extreme north and arid and desert regions. In the European part of the USSR, the northern border of the
range on the west
is

formed by the northern coast of the Kola

Peninsula (also exists on Kil'din [Island]) and eastward along the

mouths of the Kuloi and Mezen' permanent inhabitant is absent, but it is sometimes transient there. Eastwards to the Pechora delta, the border passes along the seashore, but beyond the Pechora, somewhat lowers and turns to the south, and reaches the Urals somewhat farther from the sea. However, here also there are places of transgression into the tundra (Malozemel'sk and Bol'shezemel'sk; V.Ya. Parovshchikov). In the Urals, the border falls approximately along the Arctic circle (Shvarts, Pavlinin and

White Sea coast

[rivers].

as far as the

On

the Kanin, the otter as a

Danilov, 1951).
In the northwest of western Siberia, the border passes somewhat to the north of Salekhard and along the extreme southern edge of Yamal (upper Yana river). Going to the east, it crosses Ob' gulf at its southernmost part and passes to the eastern shore of the Ob' gulf at the Arctic circle (Laptev, 1958). Farther, at the same latitude, it crosses the lower Pur and passes beyond the Taz approximately 100 km north of Sidorovsk on the Arctic circle and at the same level or a bit south of the upper Turkhan. Thence, the border suddenly rises to the north, crosses the Yenisei, and east of the Yenisei, Lomo Lake (at Norilsk) and on to the southern Taimyr, reaching there the upper Avam (right tributary of upper Pyasina) at a point a little north of 70 N. lat. It also includes the Kheta (V.N. Skalon), apparently, in its upper and middle course. The

Avam
The

is

the
is

most northernmost point of the range

a reference to otter hunting in
in the

in the

USSR.
on the
in the

otter

absent along the lower Khatanga (Middendorf, 1867).

There

is

West

Siberia,
lat.,

Gydansk Peninsula
This information
is

upper Yuribei about 7 1'* N.

if it is true,

northern part of the tundra zone (Laptev, 1958; Stroganov, 1962).

very doubtful, and

speaks only to

occasional long transgressions.

The

eastern border

is

very poorly known; however,

in

it

can be

considered established that

Yakutiya (especially

in the

west and


1299
middle) bends considerably towards the south and then again rises
strongly to the north in northeastern Siberia. This
is,

apparently,

connected with particularly low winter temperatures

of

in this part

of

Siberia, leading to formation of very robust ice cover, to freezing

many water

bodies to the bottom and even to freezing of small

the border in the eastern half of Siberia

montane creeks. The course of

in the

is

drawn

following form: from southern Taimyr, the border in a

way

not yet completely established, goes to the southeast to the upper Vilyui in the region of Chona mouth (about 6245' [N. lat.]. Then,
directing itself towards the east,
863
it crosses the Ygyatta (Igetta) and several other left tributaries of the Vilyui (Tyung, Tyukan) and passes to the lower Vilyui below Vilyuisk. Thence, the border

river

passes generally in the same direction, crosses the Lena somewhat below the mouth of the Vilyui, extends through the middle courses of the right tributaries of the Aldan the Tumara and Tompo and passes to the sources of the Indigirka and upper Oimyakon. Then, it goes along the left tributary of the Kolyma the Yasachnaya river, approximately at Sredne-Kolymsk and to the right tributaries of the Kolyma the Berezovka and Omolon (border in Yakut according to data of Maak, 1859; Tugarinov, Smirnov and Ivanov, 1934; Belyk, 1953 and Ognev, 1926, and mainly from material made available by D. Ivanov, Yakutsk).

Along
animal or
are

the described northern border (not considering the

otter is very rare

Omolon), the
its

and only individual cases where the tracks are encountered are known. The border of
lies to the south.

normal, regular occurrence

Nevertheless, there

some

less definitive data

(Maak, 1859) on occurrence (very

rare) of otter along the left tributary of the

Lena

the

Muna
little

which flows
to the south

into the

Lena from

the left at 68** N.

lat.

and, a

on the "Serka"
It is

river (apparently the Syarga) in the

upper Linda (approximately on the Arctic circle west of the Lena)

and

at Zhigansk on the Lena a bit north of the Arctic circle. These data are in poor agreement with contemporary materials

and invite doubt.

possible that they refer to occasional indi-

viduals transgressing far along the Lena. Mentions of "Alazeya in

Middle Koymsk region" (Belyk, 1953) obviously belongs in this category. The same may be said of the upper Alazeya (Stroganov, 1962, plotted this occurrence on the map very far to the north).

2
^

-5

<U

to

3 S3


1301
In the lower
it is,

Kolyma and along

the Bol'shoi and Malyi

Anyui
its

apparently, absent. Farther to the east, the border includes on

the north the

Anadyr

basin, and the range also includes

left

Belaya, Tanyurer and Konchalan (Belopol'skii, 1937; tributaries Portenko, 1941). At the mouth of the Anadyr', the border passes
to the Pacific

Ocean.
(at

The range includes Sakhalin and Shantar Islands

Bol'shoi Shantar; Ognev,

least

1929). There are no data concerning

Karagin Island. Of the Kurils, the otter was recorded for Kharimkotan (Kharumukotan) in the north and Iturup (Etorofu) and Kunashir' (Kunasir') in the south (Sergeev, 1947). On the whole, data on occurrence of otter in the Kuril chain are confused and contradictory. Several authors, both geographers (Solov'ev, 1946) and zoologists (Kuznetsov, 1949) do not refer to this species at all in the composition of the fauna, and according to new data (V.G. Voronov) it is actually absent there. The reason for mention of the otter was probably its confusion with the Kamchatka beaver (sea otter Enhydra lutris) and old information of Snou (1902) who recorded for Iturup and Kunashir the "abundance" there of "river beaver, Lutronectes whiteleyf*. The southern border of the otter's range in the European part
of the country, starting from the mouth of the Danube, generally
passes along the Black Sea coast to the mouth of the Dnepr, and
farther,

around the extreme south of the steppe

Crimea, the otter
is

to the

mouth of

the

Don.

In the

absent. In the past the otter lived,

apparently, in small creeks in the southern steppes. Farther, the

border of the range extends along the

region where the

Don and somewhere

in the

Don and Volga come

close, passes to the Volga.

Along

it,

the otter spreads to the very delta.

Don and Volga, in the Trans-Don, Kalmytsk and a considerable part of the Cis-Caucasian steppes the otter is absent. It is encountered along the Kuban', Kuma and Terek (no data concerning Manych), and the line extending along the Kuban' and Kuma represents the northern border of the Caucasian part of
To
the south of the
the range.

To

the south of

it,

the otter exists throughout the entire

is

Caucasus. The Caucasian part of the range, therefore,

isolated

from the northern. The

that, in the past,

and even the probability, exists the otter lived along the rivers flowing into the
possibility,

*This "species"

is

synonym of

lutra

Sci. Ed.

1302

Azov Sea between

864

the

steppes was not so complete.

certain

Don and Kuban', and thus isolation by the It may also be assumed that, under

populations might have and Volga. The otter is absent in the steppes between the lower Volga and Ural [riversl and in Saratov Trans-Volga. From the Volga to the
conditions,

contact between

occurred between the

Kuma

Ural river, the southern border passes, apparently, somewhere along

the latitude of Irgizi.

The

otter extends along the Ural [river] to the

[Caspian] Sea.

From
The

the

mouth of

the Ural, the border rises northward along

is entirely unclear and may be bends around the Urals [mountains]
it

the river and then turns to the east, including also the lower Ilek.
farther course of the border
It

designated only tentatively.

along the Ural river and, after rising somewhat to the north,

passes somewhere along the border between the forest and foreststeppe zone, or along the forest steppe eastward to the Irtysh, and

along

it

upstream

to its sources (recorded at

Ust'-Kamenogorsk

and Chernyi Irtysh; Sludskii, 1953). It cannot be excluded that in the cut-off section mentioned, before the Irtysh, the line of the
border passes somewhat more to the south

there

is

information

on the occurrence of otter to the southwest of Petropavlovsk and to the west of Kokchetav (Sludskii, 1953), and, perhaps, even in
the upper

Ubogan

river at

Kushmurun Lake
is

to the southeast of

Kustanai (V.G. Heptner).

Beyond

the Irtysh,

the otter

absent in the Barabinsk,

this zone, the

Kulundinsk and Cis-Altai steppes. In

range, proceeding

border of the

somewhat

to the north of

Lake Ghana, reaches

Ob' south of Novosibirsk, then goes directly south and, passing around the Altai from north [to east], reaches the Irtysh. From the Zaisan depression and southern Altai to the Pacific Ocean, the range passes to the south outside the boundary of the

USSR.
The range of
the otter in

Middle Asia

is

very narrow and

complex. Moreover,

its distribution is sporadic and it is everywhere rare. Locally, the otter is encountered only as a transient and has been observed or caught once in several years. In some regions where it occurred in the past, this animal is absent at the present time. Finally, data for Middle Asia are in general very scarce, fragmentary and in part indefinite. Therefore, it is almost

1303
impossible to outline a more or less accurate range in this region
at the

present time.
the otter must be considered as the

The Middle Asian range of

northern edge of the Near Eastern part of the general range of the
species, and in part, perhaps, the Central Asian (Pamir-Altai and
is no direct connection with the main European-Siberian region of occurrence of the species in the USSR (between the Dzhungarsk Alatau and Chernyi Irtysh otter are absent), or- it is weak and may exist in some form in the east. This is demonstrated by the systematic peculiarities of the northern and southern otters (see below). In the west, a small section of otter occurrence is occupied by the Kopet-Dag [mountains] and Atrek [river] to its mouth. This is the northern edge of the Iranian part of the range, included within the limits of the USSR from the south. Another analogous section, isolated in our country from the first, is situated along the Tedzhen river, occupying its upper and middle courses and not reaching Tedzhen city. A third section is occupied by the Kushku river,

Tien Shan, and Semirech'e). There

where, as possibly in some sections of the Tedzhen, the otter apparently occurs only as a transient from the south, and extremely
rarely (V.G. Heptner).

Concerning the occurrence of

its

otter

on the

Murgab,
though
it

at the
is

present time or in the past, positive data are absent,

not excluded, at least for

upper course.
is,

Farther to the east, the section inhabited by the otter

obvi-

ously, connected with Afghanistan, occupies the western Pamir;

the region of rivers flowing from the east to the upper Pyandzh (Rozanov, 1935), Pyandzh [itself] and the uppermost Amu-Dar'ya and the basins of the rivers flowing into it from the right, in pari.e.,

865

Vakhsh-Surkhab, Kizyl-su and smaller rivers Varzob, Khanak', Ilyak (Flerov, 1935; Chernyshev, 1958; V.G. Heptner). In the basin of Surkhandar'ya, the range includes the regions of the Tupalang and Sangardak rivers (Ishunin, 1962; V.G. Heptner); i.e. the southwestern part of the Gissar range. Along the Amur-Dar'ya, the range does not reach beyond Termez. Concerning the occurrence of the otter in the remaining part of the Amu-Dar'ya, data are absent, and apparently it does not exist along this river. The extreme turbidity of water

ticular the Surkhandar'ya, Kafirnigan,

probably prevents

its

occurrence.

Concerning all remaining parts of the Pamir-Alaisk montane system with its water

The

otter is absent in the eastern Pamir.

1304
bodies,

we have no

definite data on the occurrence of otter, exits

cept one reference to

occurrence on the Zeravshan below and

somewhat above Samarkand (Fedchenko, 1950; Ishunin, 1961). In Semirech'e, in the Tien Shan system and in the Syr-Dar'ya basin, the range of the otter occupies Dzhungarsk Altau and the
region to the north of
it

to the Alakol'

and Sasykkol' (basin of the

Kungei-Alatau
its

Tentek

river), the Karatal basin, the Hi basin and, apparently, other

(at least the

rivers flowing into [Lake] Balkhash;

northern slopes of

its

eastern half), Zailiisk-Alatau (in particular,

Chilik basin), and the

Chu

river to

lower reaches (Sludskii,

its

1938, 1939, 1953). The otter was recorded in the Syr-Dar'ya in 1948; Sludskii, 1963).

lower course and for Chatkal [range] (Zarudnyi, 1915; Kuznetsov,

Some

of the data mentioned refer only to

is,

the past, and at the present time in the otter

apparently, absent

from a series of places. For the entire remaining parts of the Tien Shan system, Fergana and adjacent places, southward as far as Surkhandar'ya and the Pamir, and westward to Samarkand there are no definite positive data on the otter's presence. There is only vague and indefinite old information about occurrence of the otter in the lower reaches of rivers flowing into Issyk-Kul' (N. Severtsov, 1873) and for Przheval'sk [city] (Kuznetsov, 1948). There are even data, that in the past the otter was distributed throughout Kirghizia (i.e., through the entire Tien Shan system, in the Alaisk range and Alaisk valley), but by the beginning of the current century, it was "almost destroyed", although, "it is possible that it was somewhere still preserved" (D. Dement'ev, 1938). Equally, there are direct references about the otter's absence in Kirghizia (Kuznetsov, 1948). At the same time, the ecological conditions permit at least sporadic occurrence of the otter almost everywhere in the mentioned mountain system.

One may

think that, in the past, the otter's natural

districts,

range occupied the whole Tien Shan and Pamiro-Alaisk

life for this

except for such places as the western Pamir where conditions of

species are unfavorable.
is

The

otter's distribution

more or

less sporadic

almost every-

where. In forest-steppe and steppe districts of the above outlined

completely absent (see for instance, Grigor'ev and Egorov, 1964). The otter nowhere occurs in great numbers and moreover, at the periphery of the range, but also in some regions far from the
northern range, there are, moreover, areas where the otter
is

1305
periphery, the animal sometimes disappears for a considerable time

and then appears anew. Local and long-distance transgressions may occur, as mentioned previously.

Geographic Range Outside the Soviet Union

This occupies
all

of Europe, to the west including Ireland, on the

north the northern part of the Scandinavian Peninsula.

On

the
is

Balearic Islands, Corsica, Sardinia, Crete and Cyprus, the otter

absent.

The range

farther includes

northwestern Africa (Algeria

and Morocco), Asia Minor, Syria, Palestine, more northern parts of Iraq except central and southeastern desert areas, Afghanistan,
Kashgaria, parts of Tibet (details unknown; apparently those con-

nected

with Himalayas),

Kashmir, Himalayas

and northern

India to the north of the Ganges. Southern India and Ceylon form
a separate region of otter occurrence. In the central parts of India,
L.

lutra does not occur,

here,

like

the

ranges
(L.

of the

and therefore, the range of this species kharza [Charronia] and thar,
nair)
is

Hemitragus) are disjunct. The northern border of the range of the

southern Indian race
/.

not accurately established. In

Central India, Lutra (Lutrogale) perspicillata lives.

Farther, the range includes

Assam, Burma, the Indochinese

Peninsula, Malacca, Sumatra and Java. In Asia, the range includes,

moreover, the Mongolian Republic (mainly the northern and montane parts of the country), northeastern China (former Manchuria), the greater part of the rest of China (except the steppe and desert regions), Japan, Taiwan and Hainan.
866

Within the limits of the outlined region of otter occurrence

outside our country, there are considerable expanses where this
species
is

absent. Thus,

it is

absent in the greater part of the deserts

of Central Asia; apparently, in the greater part of Tibet, etc. Details are,

however, unknown. In some parts of the range the otter

has been exterminated (V.H.).

Geographic Variation
Although the
though one
otter's
is

range

is

vast,
It

it

is

remarkable that
(for

its

geo-

graphic variation

very small.

is

greater in South Asia, alit

may

think that presentations of

example, Pocock,

1942; Ellerman and Morrison-Scott, 1951) are nevertheless over-

1306

Fig. 315. Species range of the otter Lutra lutra L.

V.G. Heptner.

estimated. Increase in variation along the southern edge of the

range
in the
istics

in

Asia

is

manifested both

in the

number of forms and

also

degree
of]

of their separation.
I.

Thus, for example, the character-

of Java otters, L. (L.)

barang (material of Z[oological]

in structure

M[useum

M[oscow] U[niversity])

of not only the

skull but also dentition are so considerable that,

from a purely

morphological point of view, they are analogous

acteristics.
It is

to specific char-

characteristic that all the southern "border" forms

of Asiatic otters {nair, barang, chinensis) are small, while on the

whole, geographic variation of Eurasian otters reflects Bergman's

Law

well.

certain decrease in dimensions southward

is

also

noticed within the boundary of the

USSR,

but

it

is

very weak.

1307

Himalayan
867

otters (monticola, kutab)

do not

differ in dimensions,

or differ very slightly only, from the northern otters. Moreover, in

the south, there has been observed in places lightening of general

color and of the relative color of the venter especially of neck and
throat;
i.e.,

increased contrast between the upper and lower body.

otters in the

Geographic variation of

Old World

is

still

com-

pletely insufficiently studied. This also applies to the territory of

our country. Descriptions of separate races have been done on very little material and have not subjected to serious revision. For the

USSR,
1.

the following forms

may be
I.

preliminarily accepted:

Northern

otter, L. (L.)

Intra Linnaeus, 1758 (syn. vulgaris,

baicalensis, amurensis, kamtschatica, steinegeri).

Dimensions and mass relatively larger. Color of back and sides dark-tawny, color of ventral surface relatively dark. Underfur silvery, with dark chestnut-colored hair tips. Pelage close-fitting and soft. Whole range within boundary of the USSR except the northern Caucasus, Trans-Caucasus and Middle Asia, and southern and
southeastern Kazakhstan.

Outside the

USSR in

Europe, Mongolian Republic, northto this

eastern China (former Manchuria), Japan.

The above given species description applies

1289].
It

retains

its

principal characteristics

in

the

form [p. whole huge

expanse mentioned and under all the diversity of conditions in which it exists. Thus, even the separation of the Kamchatka otter
as a separate subspecies did not receive approval. Furriers within

the outlined limits also

do not distinguish separate fur types, not-

ing only that skins obtained from Yakutiya and from the Far East
are of particularly high quality

of darker

color and have very

dense underfur. The

Eurasia
is,

entirely

weak
its

variation in otter of northern

it

apparently, associated not only with the fact that

ability to settle in

is

an aquatic form but also with

distant places.

new and

quite

2. Caucasian otter, L. (L.) /. meridionalis Ognev, 1931. Dimensions and mass relatively less large. Color of back and sides lighter than in northern form light tawny. Ventral surface also somewhat lighter. Tail monotone both above and below. Underfur silvery with yellowish tint and light

tawny hair

tips.

Pelage less close-fitting, somewhat coarse.


1308

Body

length (14)

530-710 mm;

tail

length

340-470 mm; length

3.7-8.5 kg from Okum' river basin in western Trans-Caucasus and Kura; M.P. Pavlov, 1964). In northern Caucasus and Trans-Caucasus.
(adult individuals

of hind foot 115-135

mm;

ear length 20

mm. Weight

Outside the
This

USSR in

Iran,

perhaps Iraq, Syria, Palestine and

poorly studied. In-

parts of Asia Minor.

is

a well-distinguished form, but

still

dividuals from Trans-Caucasus are quite typical, and northern

Caucasian population comprises some numbers of more darkly

colored individuals;
i.e. it

bears characters transitional to the north-

ern form. Distribution outside the borders of our country are entirely

unclear. Systematic
to the following

and geographical relationships of

this

one are unclear (see below); nevertheless, affiliation of the Trans-Caucasian otter with the southern group of forms is evident. Its range, moreover, if not completely, then perhaps to some degree may be isolated, from that of the northern

form

form by the steppes of the Cis-Caucasus. One may, apparently, consider that this isolation is greater than that from the southern
parts of the species range.
3.

Middle Asiatic

otter, L. (L.)

/.

seistanica Birula, 1912 (syn.

oxiana).

Dimensions similar to preceding form. General color is lighter back is grayish-light tawny, usually with gray hairs; sides lighter
than back, the belly very light whitish-yellow or whitish-gray.

Sometimes has yellowish tone lightening

throat forming a distintail is

guishable patch variable in size and color. The

coarse and not very close
In

bicolored
is

upper like back and lower a light dirty-yellow-white. Pelage

fitting.

Middle Asia, on northeast including Dzhungarsk Alatau; possibly also region of occurrence in Turkmenia. Outside the USSR in eastern Iran and Afghanistan. This form is well differentiated from the northern, although

among

the described type in the population are dark individuals

with fur color and characteristic very similar to that of the northern

form, especially
identical

in

the mountains.

It

is
it.

very similar

if

not
they

to the

Trans-Caucasian form

to

Together with
(its

it,

themselves represent members of the group

size not clear)

apparently of the Near Eastern type distributed along the southern

edge of the western half of the country's Asian

part.

Ranges of the


1309

Trans-Caucasian and Middle Asian form (meridionalis, seistanicd)

in their

northern section are, to a considerable degree, quite fully

isolated

from southern regions inhabited by the nominal form. This

applies to the Cis-Caucasian steppes, steppes of Kazakhstan, the

expanse between the Altai where the nominal form lives and Dzhungarsk Alatau occupied by the Middle Asiatic (Sludskii, 1953). If the otter's range in the Near East were not so limited, one could assume that forms of this type have settled (penetrated) with us from the south. The group of otters mentioned is analogous to the
southern group of races of brown bear
isabellinus-syriacus.

Systematic and geographic relationships between the mentioned

forms are not clear. They are described from quite nearby places (lower Gilmend-Tegeran) from very few materials, having similar
characteristics.
It

cannot be excluded

that, in the future,

with ac-

cumulation of new material, these forms

clear
is

may be

united. Also not

Middle Asiatic form, to the other adjacent races described from the Himalayas and northern India. The forms monticola, aurobrunnea and kutab were described from Nepal (first two forms) and from Kashmir respectively, and were also recorded in Kashmir and Punjab; i.e., in those places nearest to our southern Middle Asiatic regions in part directly
the relationship, at least of the

neighboring (Kashmir).

Leaving aside the problem of relationships among the listed Indian forms (they are hardly entirely independent), one may be assured with great likelihood that our Middle Asian (at least PamiroAlaisk and Tien Shan) otters are related to them. Apparently, they

must bear the name of the Kashmir form kutab (if it is separate from monticola, which is quite doubtful). One must also keep in view that the form seistanica was described from the delta channels of a river which disappears in the hot lowlands on the border of Iran and southern Afghanistan, while all the other named races were described from montane countries. The name seistanica was left here, therefore, conditionally, as a traditional one in our literature until direct comparison of our material with Indian, and revision of all Himalayan and northern
Indian forms. Published data (Pocock, 1941) give a completely

confused picture both of systematic and geographic relationships. It seems highly probable that otters of the Himalayas and the Middle

Asian mountains, at least to the east of Amu-Dar'ya, all belong one form, which should be designated by the name monticola. It

to
is

1310
possible that the

name

seistanica

may be

related to the group

kutab, perhaps in the form of a synonym.

Outside the boundaries of

cepted: 1) L. (L.)
/.

USSR,

the following forms are ac-

angustifrons Lataste, 1885

Algeria, Morocco
/.

(possibly a

synonym

of the nominal form); 2) L. (L.)

chinensis

869

continental China, except Tibet and northeastern China (former Manchuria), Taiwan, Hainan; monticola Hodgson, 1839 northern India (Punjab, Kumaon, Nepal, Sikkim, aurobrunnea* Hodgson, 1839 Nepal Assam); 4) most elevated Garhwal (apparently, synonym of the preceding form); kutab Schinz, 1844 Kashmir, and Tibet (probable synonym of the form monticola); nair Cuvier, 1823 southern India and Ceylon well-distinguished form); barang Cuvier, 1823 Sumatra, Java,
Gray, 1837
3)
L.

(L.)

/.

L. (L.)

/.

(the

parts),

5) L. (L.)

/.

to

6) L. (L.)

/.

F.

(a

7) L.

(L.)

/.

F.

Malacca, Indochina

(a sharply-expressed race) (V.H.).

Biology
Population. The otter belongs
animals. In the past,
affect the status of

among

number of low-density

when

exploitation did not yet considerably

of the

its stocks, the annual yield within the boundary was 5000 otters, over the course of a series of years. Annual growth of this animal ranges from 20 to 30%. Taking into

USSR

consideration the absence of overhunting at that time,

it

may be

assumed

that the

number of

otters in the

USSR

did not exceed

20,000 individuals, falling to about 10,000 in years of more intensive hunting. The smallest number of otters is in Siberia, while in West European countries, in spite of the predominance of cultivated lands and intensive settlement of the territory, it is more frequent than in the USSR. The yield from the USSR constitutes 10% share of the global one, that of West Europe a 25% share

of the skins; the cause of this

is

the fact that otter populations are

most of
bodies.

all

limited by the character of the ice regime of water

The
net,

otter

is

distributed throughout almost the

Its

whole USSR,

but very unevenly.

range

is

not continuous, but forms a sort of

all

which
of

is

generally characteristic of

animals closely associ-

ated with fresh water. Density of river distribution determines,

first
all,

density of the otter occurrence. However, there are

*Misspelled aureobrannea in Russian original

Sci.

Ed.

1311

^\":i^d

869 Fig. 316. Habitat of

otter in Altaisk preserve.

Lower Chul'chya

river.

Photograph by

G.D. Dul'keit.

Other very essential causes limiting the

number of

otters

and the
usually

density of settlement in fresh waters of the

870

USSR.
It

Habitat.
danger.

The

otter is

a true freshwater animal.

its

captures food in water, and here also takes

It

shelter in case of

usually only

comes ashore

to rest.
It

The

otter inhabits

freshwater bodies in
seashore where
it

all

landscape zones.

also lives on the

even becomes frequent locally, but usually only in the mouths of rivers, or even creeks. Height above the sea level, characteristics of relief and geographical latitude, as well as the direct influence of climate are of no importance to otter in the
presence of other favorable
intensively hunted,
it

life

conditions.

Where

the otter

is

not

does not avoid the densely populated places

or even the neighborhoods of large cities. In

are

Middle Europe, cases

known
is

of otters even living in city.

On

the whole, the

number
to

of otters
the

inversely proportional to the

number of

hunters. For

same reason, numbers of

otters usually increase parallel

increasing afforestation of a place.

1312

870 Fig. 317. Habitat of otters in "Kedrovaya Pad'" preserve, southern Primor'e. May 1963 (Photograph by A.G. Pankrat'ev).

The characteristics of river banks and their relative inacessibility humans are very important for occurrence of the otter. Almost impassable wooded banks, reed thickets and turgai, and general
to

inaccessibility of the shore zone

from land, are favorable for

it.

Because of

this, the otter

evidently often prefers forest rivers, with

greatly cluttered channels

choked with obstructions and broken

twigs and branches which

make

these rivers impassable not only

for boats, but also for a walking person. In spite of the fact that

the nature of these rivers hinders the otter itself in catching food,
it

settles willingly in

such rivers.

Rivers with transparent waters, rapid flow and rocky beds are
preferred by otters to water bodies with slow flow and polluted
water,

muddy
it

bottoms, matted water plants and opaque water

is

al-

though

also settles in such bodies. All of this

related to con-

ditions for catching food.

Abundance of

essential: of deciding significance is the

its food is also very abundance of fish in the

871

One of the main conditions in the otter's ecology also proves to be the character the water and snow regimes of the water
water body.

1313

871

Fig. 318.

Winter aspect of otter habitat on Teietsk lake

otter (here also tracks of

in Altai preserve. Tracks of kolonok and American mink). Photograph by G.D. Dul'keit.

body in the winter period. The presence of polynias [open water], thawed patches, ice holes, and "spring places" opportunities for otters to penetrate into the water and catch food there are

necessary for the

otter.

The feeding

possibilities in unfrozen sections of water bodies

and therefore open areas under ice, which are formed during winter reduction of water [level] are of no less
are usually limited,

importance

to otter.

This gives

it

the possibility of finding food

1314
under the ice without coming out onto the surface and in this way, body being utilized is extended.

the significant part of the water

With the freezing over or complete freezing of the water body, it is not suitable for the otter in winter. The impregnable ice cover and the rarity of unfrozen sections practically deny the otter the
possibility of exploiting

even those rivers richest

in food.

Pro-

longed ice cover

is

also unfavorable for these animals.

predominate

Food. Vertebrate animals, and above all fishes, everywhere in the food of the otter. Frogs, birds and mammals

sometimes play an essential, but strictly seasonal, role. The significance of invertebrates is small these are mainly large water beetles. Locally, crayfish and freshwater mollusks have some

significance.

Consumption of plant food has been noted only

for

otters held in captivity.

Volga delta, the frequency of fish in the otter's diet constitutes 88% on average for the year. In the warm period of the year, it remains at a level of 91.6-97.6%,
In the fish-rich channels of the

but in winter falls to 55.6%, while during the period

when water

body
this

are frozen,

and because of great difficulty of

all

their capture at

time

to

27%. Among

fish eaten at that time, cyprinids

constitute 60.2%, carnivorous fish

23.6%,
in

and sculpin

10.7%.
otter pre-

Fry, eaten avidly, constituting about 1/3 of

fers small fish to large. In first place

all fish.

The

frequency stands the carp

(33.8%), constituting

(24.3%);

in

25% of all fish eaten; in second place pike third roach (18%); followed next by sculpin (13.2%)

872

and shiner (11.6%). Many other fish are encountered, but in small amounts (403 samples; A.G. Romashova). In the upper Pechora, the proportion of fish in the diet constitutes about 65% annually, fluctuating seasonally from 58% (summer)
the
to

74%
i.e.

(spring).

The

significance of fish here

is

less than in

Volga

delta.

(13.4%),

Grayling (24.2%), rock sculpin (26%) and loach* rheophilic forms, predominate, constituting on the

whole

57%

bottom

79%

of fish eaten.
[fishl.

Of

these

43%
is

are pelagic species,

and
in-

Limnophilous forms constitute only 21%,

cluding only

4%

bottom [forms]. This

explained in that the otter

prefers water bodies with fast transparent water

to catch fish. Inhabitants of

where

it is

easier

deep water

whitefish,

ide, are rare

here in the otter's diet, and, as in the Volga delta, small fishes

*The Russsian word "golets" can mean

either loach or

brook trout

Sci. Ed.

1315

872 Fig. 319. Pra river

in

Oksk preserve (Ryazan

district)

cover. Path of otter (trench) along deep

trail at

habitat of otter. Complete ice -42C. January 1950. Photograph by

V.G. Heptner.

(weight to 200 gm) and fry predominate. Large fish weighing more than 1-2 kg are rarely met with; in individual cases, capture of burbots weighing up to 4 kg were recorded. Pikes are not often caught, especially small ones not more than 10 cm long. Commercial species constitute 45.5% (1530 specimens; Teplov, 1953).
In the Lapland preserve (Kola Peninsula, Kislaya river basin; 414 samples 261 excrement in summer, 38 in autumn, 115 in winter; Vladimirskaya, Lebedev and Nasimovich, 1953) fish also


1316
occupies
first

place in the otter's food. In the yearly cycle, brown

trout predominates (35.5%), followed

by pike (15.2%), burbot

(15.1%), whitefish (10.8%), grayling (8.1%),

minnow

[Phoxinus]

(7.8%), and perch (5.8%). Riverine species predominate over lake

forms and constitute 78%. Small fishes (up to 20 cm long) consti29.8%. Only 4.4% were more than 50 cm tute 70.2%, and large

The otter begins to eat large fish in the trunk region. In summer, fish constitute 23.6% of the otter's diet, in autumn 5.3%, and in winter 29.6%. The otter feeds on grass frogs mainly in winter (29.6% occurrence) when it does not have adequate food. It obtains them from the bottom of small unfrozen rivulets. In summer, it catches them along the river banks and in swamps (23.6%). It occasionally feeds
long.

on ducks
873

(teal,

golden-eye) and

mammals

(voles, water shrew,

it

squirrel) in isolated cases.

Among

insects, in winter

feeds mainly
in

on caddis

flies

(12.2%) and stoneflies (8.7%), and

it

summer

ground beetles (2.6%). Among mollusks, (1.5%) in small numbers.

In the Altai, fish constitutes

eats primarily Anadonta

food, and the larvae of caddis flies

95.9% occurrence in the otter's 83.6% (Ternovskii, 1956).

In general, the species composition of fish in the otter's food

depends on the composition of ichthiofauna of the given water body and the vulnerability of each species. In Caucasus, brown

Chondrostoma and others predominate. In forest rivers of the Volga basin, the otter feeds in winter mainly on burbot, and in summer on other bottom fishes. On the Murmansk coast, it feeds on cod, salmon trout and others. In the otter's diet, frogs are
trout, barbel,

a typical winter food, exhibiting sharp seasonal fluctuations. In the

upper Pechora, the average annual frequency of occurrence

fluctuating from
In the

is

18%,
in

10%

in spring to

26%
46.1%

in

winter (Teplov, 1953).

Volga delta

this fluctuation is

sharper

from

2.4-3.5%

the

warm

period of the year to

in winter

874 periods of

tant

food for otters

complete freezing of the water body. It in winter, but by no means a preferred food. In

and 69.2% in is a very impor-

spring, frogs are

numerous

in the

Volga

delta, but the otter hardly

catches them

at all. It

only hunts them under ice in their hibernatthe Pechora, in the

ing places where up to 1156 frogs

(A.T. Romashova).
year, the otter feeds

On

may congregate warm

per

hectare

period of the
the

more intensively on frogs (10-18%). Here,


1317

873

the otter's place of permanent Fig. 320. Polynias [open water) in Kedrovaya river residence in winter. "Kedrovaya Pad'" preserve, southern Primor'e. December 1963. Photograph by A.G. Pankrat'ev.

water bodies are less rich in fish and conditions for capturing them are, apparently, also less favorable. The significance of birds in the otter's food varies greatly. In the upper Pechora their average annual frequency is 13%. In the Volga delta, in spite of the abundance of birds, is two times less 6.1%. In the upper Pechora in summer, birds comprise 26%; among

1318

Ducks of open waters are encountered somewhat more often than those species which confine themselves to dense growths of water plants. In the Volga delta in summer, frequency of birds reaches only 1.8%, but in this
them, ducks

7.9%

and snipe

6%.

case, not ducks, but to coots and their nestlings. Cases of otters hunting moulted ducks are very rare the birds restrict themselves to places unfavorable for hunting by the otter. Apparently the

increased frequency of frogs and birds in the food of the Pechora

otter
is

associated with small numbers of fishes.

in the otter's

The mammals represented

more
rarely, other species

food are mainly water

still

voles [Arvicola], less frequently water shrew [Neomys] and

which can appear on the shore of the

across
it.

water body or

may swim

Due

to local conditions, the

significance of water voles varies. For example, in the upper

Pechora, depending on the season, the percentage of occurrence ranges from 2 to 5%, and in the Volga delta, reaches 12% both in

During the period of complete freezing, it 19.2%. then increases to become one of the major sources of food Complaints have recently appeared that the otter in the Ural [river], noticeably affects the number of muskrats (Bakeev and Koryakov,

summer and

in winter.

1960).

The

daily ration of the otter consists of about

kg of

fish.

When

constantly supplied with food, the otter eats during a night

0.4-0.9 kg of fish, in the breeding season

prolonged starvation up to 1.5 kg. (A.T. Romashova). Feeding on small fishes, the otter becomes satiated overnight with 2-3
helpings, and with a large catch

0.2-0.4 kg, and

It is

after

after

helping.

curious that

one water vole of about 145 gm can fully satiate it (A.T. Romashova). This is probably conditioned by the higher nutritional value of warm-blooded animals in comparison to fishes. Home range. The home range of the otter is limited in width not wider than 100 m on each to the narrow [river] bank zone

bank. Depending on the food capacity of the water body, the home range may be unitary or consisting of a series of separate ranges,
separated from each other by non-feeding areas.
section and
its

The length of
sections

the

area depends also on the abundance of food and

capture.

conditions for

its

The separate feeding

may form

a chain extending for 15-20

the otter
ies,

km

along the course of one river, or

may

periodically visit a series of adjacent lateral tributar-

reaching them by way of their mouths or crossing the

1319
watersheds on land. In water bodies with abundant food, the otter
leads a settled life in a section extending from 2 to 3-5 km. In the
great taiga rivers of the Pechora basin, the average length of the

home range

for a series of years

fore, the area of the

was 6 km (Teplov, 1953). Thererange inhabited by the otter fluctuates from 4-

On the Kola Peninsula, the otter covers 4-6 km 24 h period and sometimes more (Vladimirskaya, Lebedev and Nasimovich, 1952). In winter, when roaming around the watersheds, and also with
12 to 300 hectares.
in

875

moves along snow and Kola Peninsula, leading almost a migratory mode of life in winter, the otter commits a transgression into a neighboring section 20-30 km in length and sometimes including neighboring rivers and creeks. On ice, it moves up to 15 km, and on snow, up to 8 km per 24 h period. Burrows and shelters. Within the boundary of the home range, the otter usually has a permanent burrow, and also a series of temporary shelters and hiding places. The latter are entirely variable depending on local conditions. These may be eroded areas of steep banks protected by tree-roots, heaps of driftwood and fallen branches, and caves in the rocky shore, etc. The permanent burrow is often constructed in a steep, although
strong freezing of water bodies, the otter
river ice

15-20

km

per 24 h period.

On

the

not high, bank.

Its

entrance opens under water, at a depth of 0.50-

The sloping passage, 1.5-2 m long and about 20 cm in diameter, leads to a nest chamber 0.4-0.5 m in diameter, often lined with dry grass or moss. In one case, it was found at a depth
0.75 m.

of 20

cm from

the soil surface (Teplov, 1953). In the family buris

row, the nest chamber

always located so that

it

may be

sub-

merged only during spring flooding. During that period the otter, like the beaver and desman, is deprived of permanent shelters. In low-lying places where high level of groundwater does not permit
construction of burrows, the otter builds a shelter for the
litter

of

young

in

high piles of reeds or fallen branches,

silt,

in the

heaps of
young,
a
lair.

driftwood half-covered by sand and

In well-protected places, the otter

flotsam,* and other cover.

birth to
its

sometimes gives

even on the surface, but does so

in a carefully

concealed nest

*In the Russian original this

of "splavnik" (flotsam)

word

is

misspelled "plavnik"

(fin, flipper)

instead

Sci. Ed.

1320
In winter, in

Anadyr

Territory, the otter

a shelter under a steep overhanging

sometimes settles in bank which has an exit to the

sub-ice space.

Its activity is

thus hidden to a considerable degree

from the eyes of observers. On Kola Peninsula, otter forms occur under snow, rarely on the shore or on snow-covered rocks. "The dining room" is located on rocks which have on one side an edge sloping to the water. "Latrines" occur under spruces with the crown
hanging near ground, usually directly on ice (Vladimirskaya, Lebedev and Nasimovich, 1953).

875

Fig.

321. Shore rocks with niches and clefts

entire

shelters of otter in the course of

the

year

"Kedrovaya Pad' " preserve, southern Primor'e. Photograph by A.G. Pankrat'ev.

May

1963.

1321

Daily activity and behavior. The otter leads mainly a crepuscular-nocturnal

mode

of
it

life.
is

It

is

especially active on moonlit

it

nights. In places

where

not disturbed,

sometimes also apin

pears during the daytime; this

more often occurs

winter than in

876

summer. Increase in diurnal activity in winter sometimes manifests dependence on diminished conditions of illumination under the layer of snow and ice, which hinders the otter in catching fish at

Fig. 322. Winter shelter of the otter on an eroded bank. A bunch of dry grass at the edge of an ice-free area, which the otter was carrying to the shelter, and dropped upon the approach of a human. "Kedrovaya Pad'" preserve, southern Primor'e.

January

1961.

Photograph by A.G. Pankrat'ev.

1322
night under the ice (Teplov, 1953).

On

active round the clock (Vladimirskaya,

the Kola Peninsula, it is Lebedev and Nasimovich,

1953).

The
for
its

otter leads a secretive

mode

of
it.

life.

It

hunts, by waiting

prey on shore, or by pursuing

silty

Obstructions in the water

body, a

bottom, unclear water and growth of water vegetation

otter,

hinder capture of food by the

but at the

same time increase

the protected nature and safety of the habitat.

On
more.

dry land, the otter

is

clumsy and

less mobile, but has en-

durance, and even in deep snow, progresses for ten kilometers or

No dog

at all

can overtake the

otter,

even following fresh

tracks. In water, the otter is very quick, dexterous

and mobile.

It

catches large fish, by diving under them, seizing the abdomen, or

rarely taking the back.
It

may happen

that a very large fish pulls

away from its teeth and escapes with wounds (A.T. Romashova). Diving under the water, the otter is at first not evident, but then its path may be followed by the bubbles exhaled when it breathes out.
For breathing, when there is danger, the otter sometimes thrusts only the tip of its nose to the surface. Sallying out of the otter onto
shore most often occurs in definite, well-concealed places, and
usually they have the character of beaten tracks, similar to those of
the beaver. In

some

places, so-called "otter slides"* occur.

are smoothly polished, well-beaten paths

These on steep slopes of the

bank, often

damp and slippery. There are observations that otters "amuse themselves" by sliding many successive times along these
steep tracks into the water.

Moving along
a running

the surface of the ice,

jump, slides along on its belly, leaving a long trough-like track. Such behavior in the otter, as in mink, is explained by the need to dry the wet fur (Ternovskii,
the otter quite often,

making

1956).

The
ming.
It

otter

mainly hunts "stationary" fishes or those slowly swim-

also hunts at "fishing holes" and hunts fry on small bars.

When
877

hunting, the otter

swims with
its

its
it

head and part of

dives.

its

back

on the surface. After seeing

stay together for the

prey,

However, females and their litter whole autumn and winter, and usually remain with it until mating again. The female defends and protects the litter, and the male, apparently, sometimes participates in its upOtters usually live singly.

bringing.
*Literally, "sliding hills" in Russian original

Sci. Ed.

1323

Seasonal migrations and transgressions. The otter

tremely mobile animal. However,
its

is

an ex-

frequent wanderings, from

one water body to another, giving the impression that it is migratory, nomadic animal, actually take place within its home range. These wanderings are explained by water bodies becoming fooddeficient especially in winter, with an unfavorable ice regime, the

home range

occupies a considerable length of shoreline. Neither

can the otter's crossing of watersheds be called migration. These

also bear the character of

movements within

a vast

home

range.

Reproduction. Reproduction in otters

studied.

is still

very insufficiently
exist.

series of contradictory points of

view

These, as
to the

well as the existing facts,

existing level of our

may be harmonized according

in the

knowledge

following way.
is

1.

As

in

many
is

species of the mustelid family, there

in the river otter.
is

a latent period in

embryonic development

The duration of this period

is

completely unstable and

determined by environmental factors.

liable to indi-

In several sections of the range, the latent period

vidual fluctuations and in others

at the

it is

more or

less stable

same time,

different in different sections. After the

and is, end of

pregnancy apparently proceeds for about 60 days with2. Corresponding to variations in duration of the latent period, the times of estrus and mating are subject to some
this period,

out deviation.

deviations.

Most

frequently, apparently, estrus and mating take place

last

during March and

delta region (A.T.

about one month. This applies to the Volga Romashova), Mari Republic (V.A. Popov), upper Pechora (Teplov, 1953) and southwestern Kalinin district (P.B. Yurgenson). The date of beginning of this period is changeable
(February-April).

On
the

the other hand, there are reports of otter cubs of one and

same age (one month) found in June and October-December. Quite well-developed embryos were found in otters caught in January. In February, an otter cub the size of a domestic cat was taken. Such dimensions of young were attained both in January and July. An otter cub which could hardly walk was once found on the snow (A.T. Romashov). The data obtained from the tributaries of Amur
also confirm that otters reproduce at any time of the year

"like

dogs". There

is

a report that in the Caucasus, estrus in otters oc-

curs at the beginning of winter. For the Volga delta, the possibility

of mating in times other than

March

is

not excluded, but such

exceptions are rare. Late

litters

usually die in winter. Parturition


1324
occurs here in June-beginning of July. In the upper Pechora,

it

occurs at the end of April-beginning of May, and in June active

young

otters are observed.

On

18 June, two young males were

caught, with the eyes opened, but the teeth had barely erupted

body length 33.5 cm; V.P. Teplov). Most often, May. Previously mentioned differences in dates serve as adaptations to the local ecological conditions. The number of young in the litter varies from two to four. Individual cases are known when there were 5 otter cubs in a litter (Khorol river, Ukrainian SSR). In the upper Pechora, no more than two otter cubs were observed in litters. The number of young is closely related to the degree to which environmental conditions are favorable. In England, the newborn cubs are encountered at any time of the
(weight
1

kg,

births occur in

year, not excluding winter (Stevens, 1954). In

one region the majority

of births took place late in autumn and in winter (October-February),

and

in others

in the

period from January-February to April inclu-

sive. In

134 cases (for several years), the occurrence of new-boms

as follows: January

was distributed by months of the year

878

14;

April 10; July 11; October 12; February 8; May 11; August 13; November 11; March 10; June 11; September 12. 11; December

In the British Isles, winter is mild,

and the

otter is not subject

there to sharp seasonal fluctuations in conditions of existence. In

is repeated each month until mating accompanied by a sharp whistle produced by males. At this time, a fight may occur between them. Pregnancy lasts 61-63 days. Duration of the latent period is variable and may be more than 9 months.

captivity, estrus in the female

is

productive. Mating

is

tary.

Growth, development, and molt. Data are few and fragmenYoung animals are born blind and without teeth, with short
at the

dark underfur. They begin to see

age of about 35 days

(Stephens, 1954). In the female, there are three pairs of nipples;

however, usually only a part of them functions. Young females gain in dimensions and weight more slowly than males. Young
otters gain about

400 gm in weight each month, and at 10 months weigh about 4 kg. At the age of one year, they attain the size (but
is

not the weight) of adult individuals. Sexual maturity in individual cases (in captivity)
attained in the second year, and probably

more

often, in the third.

1325

Fig. 323. Otter

on the

ice of a creek near an otter slide.

southern

Primor'e.

January

1961.

"Kedrovaya Pad'" preserve, Photograph by A.G. Pankrat'ev.

is

Molt

in the otter

proceeds very gradually, and

little

noted.

lation dynamics. Other than

Enemies, diseases, parasites, mortality, competitors, and popuhumans, the otter has no dangerous
it

enemies. Neither has

them.

any serious competitors.

its

It

usually forces

out mink from the borders of

home

range, often destroying

to as

On

the

Kola Peninsula, the wolverine was referred

an

enemy of
is

the otter, and the white-tailed eagle as a competitor

(Vladimirskaya, Lebedev and Nasimovich, 1953). Beaver activity

usually useful for the otter

in

winter the latter uses beaver

outlets

from under the ice. Diseases of epizootic character are unknown. In captivity, coccidiosis infection has been recorded. In
pulmonary tuberculosis and endocarditis were diagnosed (Stephens, 1954). Helminth infections are not major. The main cause of mortality, especially up to one year of age,
is

879

a severely emaciated otter killed while attacking a domestic goose,

unfavorable weather conditions in winter, in particular, the ice

regime. Late litters usually die in winter. In the upper Pechora, a connection exists between the survival of young, magnitude of

autumn flood favoring the formation of sub-ice spaces and the

1326

879

Fig. 324.

Track and prints of night

18

life

of an otter family

female with young.

December 1964. Photograph by A.G. Pankrat'ev.

duration of the ice-cover period.

The

otter population there is

determined by intensity of reproduction and survival of young. The average percentage of young in otter populations was 19.0,
with variation from 12 to 26%,
1953,
1954).
otter
i.e.

more than 2 times (Teplov,

The

responds very quickly and positively to prohibition

it.

The cultivated landscape does not directly prevent the growth of the otter
of hunting (launching)* with real observance of

population
tion.

if

they are protected from direct pursuit and destrucrise

Only measures giving

to

impoverishment of the

ichthyofauna of water bodies have proven to be harmful: floating

forest litter,

sewage drainage from

difficult to

industrial enterprises, etc.

Field characteristics. Sightings of river otter are rarely successful.

otter,

and

of

this

confuse with any other animal. Tracks of excrement are often encountered. The large tracks animal most often bear well-marked outlines of the swimming
It is

their

Meaning

of parenthetical word "zapusk", or "launching" not clear

Sci. Ed.

1327

membrane between
that of the

the digits. Its track

it

is

clearly separable

from

beaver

in that

usually extends along the bank at the

edge of the water, while the path of the beaver is always perpendicular to the line of the bank these are its sallies onto the bank. Excrement of the river otter usually possesses a greenish tint,

and
880

is

often watery.

The remains of
them

fish

bones and scales of

it

fish

are typical of them.

From mink excrement

[feces]

differs in large size.

The

otter usually leaves

water body or on the bank, as well as

often buried.
Its

on rocks in and around the in holes on sandbars. It is

it.

urine typically has a strong odor peculiar only to

its
it

On
with
its

snow, the otter leaves not only

its

footprints, but also the

track of

dragged

tail.

In deep

snow

makes

whole furrow

body.
is

On

river ice, a long ribbon of a trail (sliding after

running)

typical, as are tracks

emerging from water.

snow for 10-15 cm; the length of its leaps equal 50-110 cm more often 60-80 cm; the length of the trail made sliding on snow is 2-3 meters (Vladimirskaya, Lebedeve and Nasimovich, 1953). The otter sometimes proceeds for up to 6 km along ski-tracks. The weight loading on 1 cm^ of the foot
The
otter sinks into the

surface

is

equal to 53.2

gm

for the

male and 37.4

gm

for the

female (Teplov, 1953). In the period of reproduction, the otter's

presence
881

may be

revealed by whistle of males at night (P.Yu.).

Practical Significance

The

river otter provides very valuable fur

beautiful,

warm

and durable. From it are manufactured fur articles in natural shape but with plucked guard hairs, or even dyed a black color as

Such articles are usually sold as a "true seal". The otter is not numerous and is captured in limited numbers, more or less occasionally and accidentally. At the present time, capture of this species is carried out by licensed trappers. In RSFSR, 6100-6700 otters were trapped by license in 1956 and 1957. The annual catch for the USSR was about 7.5 thousand for the same years. The world catch of otter before the Fatherland War [World War II] was 80-100 thousands. Due to the small number of otters in the USSR, the harm from it to the fishing economy is negligible. The otter is usually hunted with guns and dogs, more rarely by still-hunting on moonlit nights. Harvest by jaw-traps is of a certain
"sealskin".

1328

880

Fig.

325. Tracks of an old otter male on light, wet new-fallen

"

"Kedrovaya Pad'

preserve,

southern

Primor'e.

snow on the ice. March 1962. Photograph by

A.G. Ponkrat'ev.

significance, but
able. All other

882

lowed as The otter is easily tamed and capable of domestication, and after a more detailed study of its reproductive processes might be bred in captivity. This is, however, possible only where there exists abundant and cheap food (fish) (P.Yu.).

good traps for catching this animal are not availmethods are either not productive or are not alinhumane.

1329

;''

v#

It, e"i

.^<!4.^i
'

'i''

0.

;i

326. Prints of hind and fore feet of otter on a sand bank and the scheme of arrangement of the tracks during different gaits. Neya river, Kostromsk district. 24. X-1950. Sketch by A.N. Formozov, about 2/3 natural size.
Fig.

1330

Genus of Kamchatka Beavers or Kalans* Genus Enhydra Flemming, 1822

1816. Pusa. Oken. Lehrb.
d.

Naturgesch.,

3, p.

985.

Nom.

praeocc.

Pusa Scopoli, 1777, Phocidae. 1822. Enhydra. Flemming. Philos. of Zool.

Linnaeus, 1758.
1827. Latax.

2, p. 187.

Mustela

lutris

Gloger.
p.

Nova
511.

acta phys.-med. Acad. caes. Car.-

Renaming oi Enhydra Flemming, 1822. 1829. Enhydris, Fischer. Synops. Mammal., p. 228. Substitute for Enhydra Flemming. Nom. praeocc. Enhydris Merrem, 1820,
Leopold, 13,
Reptilia (V.H.).

Measurements very large largest in the subfamily and family. General body construction heavy, trunk very long, cylindrical in form. Limbs short, hind [limbs] displaced backward. Foot flipper-like, paw of fore limb short, digits unsegmented externally.
Tail muscular,
its

length constitutes one-quarter to one-third that

of body.

Fur long and very dense. Color tawny and brown

ternally, glandular areas at base of tail absent.

in tone.

Ex-

Two

pairs of teats,

but only one functions in female.

Skull massive and heavy.
It

is

high and broad, with facial

portion strongly shortened and elevated anteriorly, and terminated

nearly vertically. Postorbital constriction only slightly narrower

than interorbital. Brain case short, but very wide. Mastoid width

only slightly less than zygomatic. Protuberances and crests of skull

sharply defined; not only occipital but also sagittal crests well

developed.

Dental formula

C-P M = 32,
2

Incisor formula 3/2

13 13

i.e.

somewhat reduced.
carnivores (one

rare exception

among

bear has 2/3).

form of anomalous deviation, an additional underdeveloped lower incisor may be manifested. It is probable
a

As

that in

milk dentition

is

*"Kamchatka beaver" was

traders.

the

At present, the accepted

in this

common name originally bestowed by Russian fur common name in English is "sea otter," and it will
name "kalan"

be used

account, together with the native

Sci. Ed.


1331 Anterior premolars with obtuse apices, posterior premolars

and molars are almost flat, with low, blunt, rounded apices. "Carnassial" structure of upper third premolar not expressed. It has a rounded triangular form, with one angle directed inwards. Upper molar considerably larger than last premolar, of rounded triangular form, one side directed forwards toward premolar and other side inwards. First premolar only slightly shifted deeper into toothrow. Lower carnassial tooth not developed either. This is an extreme
deviation in the family in the sense of loss of "carnassial" structure

of the dentition formula,

(Lutra). Here, the cutting

is

also observed in badgers and otters

is

form of the tooth

entirely lost.

Such

a structure itself represents an adaptation to feeding on sea-urchins

and mollusks.
883

Vertebrae 50-51 (C
region elongated and
articulate with sternum.

7,

14,

6,

3,

Ca 20-21),

thoracic

composed of 14

pairs of ribs, of

which 10

Scapula broad with poorly developed acromian process; pelvis narrow and long, situated to a considerable degree parallel to vertebral column, humerus very massive,
noticeably spirally twisted.

Femur

shortened.

Stomach relatively large, intestine very long (exceeding body length by approximately 10 times), gall bladder present. Ligamentum teres absent. Lungs relatively large, diaphragm lies obliquely: fixed more posteriorly on dorsal side than on ventral (characteristic
feature of truly aquatic animals, associated with increased dimen-

sions of lungs).
Testis located under skin (scrotum
is

not formed).

On

the whole, in

all

of

its

morphological features, the genus

Enhydra represents an extreme degree of specialization in the sense of adaptation to water, not only in the subfamily and family, but also in the order as a whole. From a purely biological and bio-morphological viewpoint, sea otters occupy an intermediate position between those types of aquatic mammals which are represented by pinnipeds and terrestrial mammals, standing in the series relatively closer to the former. Of course, the described genus must not be considered as genetically linked between these groups; however,
it

indicates one of the possible biological stages in the path

of transformation of terrestrial

mammals

to aquatic.

purely marine

and in part and marine invertebrates (mainly sea-urchins).

form of temperate and cold temperate, warm temperate waters, specialized for feeding on fish
littoral

1332
Distribution restricted to shores of the northern part of the
Pacific Ocean, north
the west.

from 23 N.

lat. in

the east and 40 N.

lat. in

The genus Enhydra is restricted very sharply, and stands apart from other genera of the subfamily, especially Lutra and Pteronura. In fact, differences from the genera Aonyx and Paraonyx are also just as sharp, and attempts to relate them are deprived of any basis. On the whole, the genus Enhydra contrasts with the remaining genera of the subfamily, each of which is nearer to one another than to Enhydra. It represents an extreme type, and a terminal link in the development and specialization of the otter group. Attempts to separate it even into a special subfamily have been made more
than once.

The

origin of the genus

is

usually associated with the extinct

genus Potamotherium comprising several species (at least two) from the upper Oligocene and the lower Miocene of Europe. Connecting links between Potamotherium and Enhydra are not clear;
in the

mind of some authors

(Pohle, 1919), for this

may

serve one

of the fossil European species of the present African genus Aonyx

{A.

hessica); however, this

is

doubtful.

known from
species
otter].
is
It

the upper Pliocene,

when

it

The genus Enhydra is was represented by the

extinct species, E. reevei, in the North Atlantic (England). This

considered the direct ancestor of the present kalan [sea

is

supposed

Enhydra colonized
possible that the

from the Atlantic, Ocean (Pohle, 1919). It is entirely path of colonization was the sea washing norththat in the Pleistocene,

the Pacific

ern Eurasia. Generally, the geological history of the genus

clear,

is

un-

and there
is

is

a basis for

assuming

its

earlier occurrence in

the Pacific Ocean.

one of the characteristic species of the northern Pacific ("Beringian") center of development of aquatic and inshore fauna, in which are included such remarkable forms as Steller's sea-cow,
This
sivach [Steller's sea-lion], and a series of bird species (black guillemots, cormorants).

The genus

includes only one species: Enhydra lutris Linnaeus, 1758.

Its

One
full

of the most valuable fur-bearing species.

its

falls

under

protection throughout

entire range (V.H.).

1333
884

KAMCHATKA OR SEA BEAVER

Enhydra
1758. Mustela
lutris.

[OTTER],

OR KALAN'

lutris,

Linnaeus, 1758
1, p.

Linnaeus. Syst. Nat., Ed. X,

45.

ComNorth
allg.

mander islands^. 1777. Lutra marina Erxleben.

Asia and America.

Syst. regni. anim., p. 445.

1799. (1800) Lutra gracillis. Bechstein.

Thomas Pennants

Uebers

d. vierfuss

Thiere, 2, p. 177. "Statenland"

one of
986.

the southern islands of Kuril chain, apparently Kunashir.

1816. Pusa orientalis. Oken. Lehrb. 1827. Lutra

stelleri.

d.

Naturgesch.,
p.

3, p.

Lesson. Man. de Mammal.,

350.

156. Kamchatka.

1922. Enhydra lutris kamtschatica. Dybowski. Arch. Tow. Nauk.

Lwow,

1,

p.

Nom. nudum
Diagnosis

(V.H.).

Only species of

the genus.

'The true Russian name of this species must be considered "Kamchatka bobr" (more correctly bobior), or "sea bobr". The old popular name of the Bering Sea "Bobr Sea" was derived from this word. The name of the animal came into the

language of literature through Pushkin

("Its

bobr collar silvers as the frosty dust").

the

The word "kalan" somewhat modified from

word "kalakha" was used by Russian people only in the 18th century and then very rarely (Pallas, 1811). At the present time, because of the danger of mixing it up with the river bobr [beaver] (!), some
zoologists use
it

in

our literature. Following this "principle", evidently such names as

lily)

"sea slon" (northern elephant seal) and "sea lev" (sea lion), "sea ezh" (sea urchin),

"sea liliya" (sea

there.

etc.

must also be changed. The name "kalan" came into

It

daily use, but must

now

be considered as artificial and bookish.

is

not

known

the young animal suckling still feeding on milk "medvedok" [young cub], the semiadult (1 year) "koshlak", the female "matka" [dam], and the adult male "bobior, bobr" (=otter). ^The assignment of Linnaeus "habitat in Asia et America septentrionali" was fixed in that sense by Barabasch-Nikiforov (1947). Some authors consider Kamchatka as the type locality of the species (Ognev, 1931; Ellermann and Morrison-Scott, 1951, and others), which has little basis. To avoid undesirable nomenclature changes (renaming), the submitted interpretation of the restricted type locality must be sustained. Steller's description of "Lutra marina" was given before 1758 (in 1751) and has no nomenclatral significance. "Viverra aterrina" of Pallas is sometimes submitted in lists of synonyms (Yakobi, 1938) but has no relation to the described species (Heptner, 1934). It is a kharza (see [Vol. II, pt. lb]). In Erxleben, the name L. marina is associated with the given species only partially, because the author also had in mind
the Brazilian otter.

name Fur manufacturers name

Such
is

also the

"sea vydra" [sea otter] a translation from German.


1334
Description^
Externally the sea otter
thicker than head, and
posterior.
to the highest

is

degree unique. Trunk

elongated, of cylindrical form, and quite massive.

short.

Neck thick Limbs very short, hind ones displaced Animal appears clumsy and heavy appearance. This
is

impression
very long
in

partially

due

to not

only very luxuriant, although not

fur,

but also to skin loosely enveloping the trunk

land,

as if

a sack.

On

movement of
is

the animal
it

is

hesitating and
its

relatively slow; while lying or

moving,

usually flexes

back

strongly. In water,

it

very agile, mobile and flexible.

vibrissae and small eyes. Ear

Head rounded, with dense long

slit-like,

pinna small (resembles external ear of seals), auditory openings

may

close (during submergence into water), nostrils are

slit-like, closeable.

The

iris is

blackish-tawny.

885

Foot flipper-like,

all

digits invested with

common

covering

and fused

to last phalanx; digit

(outer) very long, remaining

gradually decrease in length to very shortest first; foot above and below covered with wool. Fore limb strongly shortened. Paw short, digits not segmented externally and only weakly defined. On lower side is found somewhat bulbous, round bare area, in anterior part of which four small parts marked by slight grooves corresponding one common narrow part extending to digits, and in posterior

transversely, [Fig. 328].

second external part of fore limb

in-

cludes two very long digits (on upper surface, there are corre-

each.

spondingly two claws), remaining one of three include one digit Claws of both fore and hind limbs very small, dark horn in

color. Tail covered with a dense short fur, relatively short (about

25-30%

of the body length), muscular and slightly flattened

dorsoventrally.

Fur of sea otter not especially long, but exceptionally dense, and silky. It is one of the most precious furs, in fact is not comparable to any other fur as regards its beauty (especially "gray
soft
otter")

and durability.

Guard hair and underfur differ little from each other in length, and are, moreover of more or less equal length throughout the whole body. The longest fur is in the middle of the back. In

Commander
'Mainly
Z[oological]

otters,

average length of guard hairs here

to

is

27.7

mm,
of

according

Barabash-Nikiforov
U[niversity].

(1947)

and

materials

M[useum

of]

M[oscow]

1335

Fig.

327. Sea otter, or kalan, Enhydra lutris L. Sketch by A.N. Komarov.

Fig.

328.

Lower surface of

fore

foot of kalan, Enhydra N.N. Kondakov.

lutris

L.

Sketch by


1336
of underfur
tively;

22.5 mm. For withers, 24.3 and 19.5 mm, respecfor sacrum 24.5 and 21.7 mm. On scapula, hair length

is

886

mm; on middle of side, 25.9 and 20.3 mm; on hip 20.3 and 16.4 mm. On breast, guard hairs have length of 25.3 mm; underfur 15.0 mm; on belly, 23.4 and 17.0 mm; on groin, 21.1 and 15.0 mm. Maximum thickness of guard hairs is on breast (169.5 micron). On belly, it equals 147.6 [|J.]; on middle of side, 136.6
24.5 and 19.0

on sacrum, 125.0 [|i]; on middle of back, 123.9 [|i]; and on hip, 115.3 [|x]. Tail is covered with shorter hair compared to trunk along its dorsum, length of guard hairs and underfur is 19.0 and
[\i];

16.5
hairs

mm;

along venter, 18.5 and 15.1

tail

mm

[respectively].

Guard

even coarser than on whole trunk (except breast): their thickness is 153.2 [jx]. On lower surface of tail, their thickness is 115.7 [|i] (all figures are mean of 50 hairs). On 10

on dorsum of

mm^

of back, an average of 12.4 (in summer), and 14.6 (in winter)

guard hairs occur, as against 1910 and 1725 underfur hairs, respectively; on belly in summer, there are 20.2 and 1674, and in winter,
17.2 and 2221, respectively (Barabash-Nikiforov, 1947).

General color of fur is dark brown. Facial portion of head brownish-gray, light straw or almost white; vibrissae are white.

886

Fig.

329.

Kalan on Mednyi Island, March 1964. Photograph by S.V. Marakov


1337

Rear portion of head and occiput dark brown, sometimes mixed with separate white ("gray") and light reddish hairs (guard hairs with light tips). Fur on neck, withers and back dark brown, of an especially dense, deep, almost black, color on back. On the neck and withers, there is a significant mixture of yellowish-gray hairs, white and reddish guard hairs, and also on base of tail. Lower surface of neck and breast brownish-gray or even dirty-white. Abdomen dark brown, only a little lighter in tone than back. Transition in color from back to abdomen is gradual. Fore and hind legs
similar in color to sides, but slightly darker.

Gray

hairs are absent

on them. Upper side of

as in

tail

same color

as

back and lower side

kalan, chiefly after

abdomen

(prevailing type of

Commander

Barabash-Nikiforov, 1947).
Individual variation in color
is

significant. General tone of fur,

from the [above-]described and most widely-distributed type, proceeds through gradual gradations to deep pitch-black, or through

Fig.

330. Old kalan.

Mednyi

Island,

May

1960.

Photograph by S.V. Marakov.


1338

tawny

to light

brownish. These differences are determined by

a

characteristics of pigmentation of the terminal part of the hair

their bases

have

very constant gray-straw color. Not rarely,

there are significant exceptions to the described lightening of the

887

anterior part of the trunk, and individuals are

sometimes encoun-

tered with an almost white (cream-colored) head, neck (upper and

lower) and breast. Otters with white heads and dark-sandy trunks

belong

to the category of

otters are also

extreme color deviants; completely white met with (albinos; Fig. 331).
is

Admixture of gray hairs

entirely variable. Usually, they are

distributed on withers and anterior part of upper body, but

some-

times occupy a greater expanse. Sometimes they are unevenly distributed, as in patches of different degrees of density. In a series

of cases, they
tion, the path

may be completely

absent. Beside individual varia-

and condition of molt itself have influence on degree of gray hair development (see below). Seasonal changes in color and density of fur are insignificant. As in several other typical aquatic animals, molt in sea otters is not
limited to a defined period, extending throughout the entire year,
is always actually complete, or almost complete. However, molt is more intensive in summer months, which is responsible for noticeable differences between "summer" and "winter" fur. Before shedding, there is, in places, lightening of guard hairs, and therefore during the period of more intensive molt, the quantity of light (gray) guard hairs on the skin is greater. Therefore, intensity of molting, i.e. mainly summer animals, appear "gray". Except change in quantity of gray hair, which gives lighter general fur color its tint, the basic color of the animal also changes

proceeding slowly, and pelage

somewhat. In summer, it acquires a lighter brown tone, apparently because of fading of the fur under the influence of the sun's rays (Reshetkin and Shidlovskaya, 1947).
Sexual differences in color are absent, or
hairs
is

only revealed in

slightly lighter color of females. In females, the quantity of

is,

guard

on the whole, somewhat

less, in

connection with which

their fur is generally

888

somewhat

softer.

Age

variations in color are quite significant, and proceed slowly.

Recently-born animals (medvedoks) have light brownish-red or reddish-tawny fur. Head and neck are lighter than remaining part
of trunk, but the contrast
is

not as sharp as in adults.

fur
is

Abdomen

is

somewhat darker than back;

dense, but relatively short, with

1339

Fig.

331.

ground

Albino kalan.

group of kalans

in

On

rocks, glaucous

Zapalat Bay on Mednyi Island. In the forewinged gulls {Larus glaucescens Naum.).

6 August 1962. Photograph by S.V. Marakov.

coarse guard hairs and

in

is

quite harsh to the touch.

less. In a yearling,

Such

fur occurs

animals 5-6 months old or

contains

but not yet adult,

otter (koshlaka), fur

does not differ in color from that of adults,

but, apparently,

still

some quantity of coarse guard pelage

of the medvedoks. In both of these stages, gray hairs are absent.

Maturity

is

attained
iris is

by sea

otters, apparently, in the fourth year. In

young, the

nut-brown.
in color are

Geographic variations

poorly studied and, gener-

ally speaking, are insignificant.

Skull large and massive, high and broad, but short, with sharply

shortened, high facial portion, as

332].
Its

if

vertically cut anteriorly [Fig.

length

is

approximately two times less than cranial part of


1340

Fig.

332. Skull of sea otter, or kalan, Enhydra lutris L.

skull.

Upper

profile of skull almost horizontal,

posteriorly. Cranial portion of skull relatively very large

its width only slightly less Zygomatic arches massive and strong widely separated

weakly rising and wide than width of zygomatic arches.

laterally.

Auditory bullae large, located transversely, auditory openings very

small, directed obliquely upwards. Nasal bones short and broad.

Upper processes of premaxillae narrow, extend

to nasal bones, but

contact only their anterolateral processes. Postorbital processes

1341

weakly developed. Postorbital constriction short and broad,

constituting about one-fourth condylobasal length, but less than
interorbital constriction.

Nasal openings terminate almost vertically, very wide their width greater than width of orbit (from postorbital process to zygomatic arch). Nasal turbinates very strongly developed intricately convoluted, filling the whole nasal opening and extending
its outer termination. Hard palate cleft behind cheek teeth very short and almost unconstricted. Sphenopalatine notch very short and wide. Diameter of infraorbital foramen is nearly equal to diameter of alveolus of upper canine. Mastoid proc-

forwards nearly to

esses isolated, blunt and massive, directed obliquely downwards.

Opening of jugular foramen very large

cipital processes

its

diameter approximately

equal to diameter of alveolus upper canine or slightly less. Paroc-

very weakly developed. Coronoid process of lower

mandible very large (60-80% of mandible length from angular

890 process to alveoli of incisors), angular process small.

Protuberances

and crests of skull on the whole, and

Dental formula

sagittal occipital crests in

particular, sharply expressed in old [individuals].

see the

characteristics of the genus.

Individual variations in skull lie in several changes in general

dimensions and proportional

there
is

details. It is a fact, that not rarely

in places

asymmetry of the

skull

somewhat

stronger

development of the left side of the braincase. This phenomenon is observed not only in old animals, where it is manifested in particular in twisting of the sagittal crest, but also in much younger
animals.

Age changes

in skull

very great. In young, postorbital proc-

esses weakly developed, width of postorbital constriction less than

width of interorbital area; facial portion of skull relatively still shorter; braincase, on the contrary, still more developed, more

rounded and swollen; crests (sagittal and occipital) absent; suture between basioccipital and basisphenoid not fused*. In the upper toothrow, last tooth is third permanent premolar.
Sexual differences
relatively
in

skull not sharp, but obvious. Skull of

is

female, beside smaller general dimensions,

distinguished by

its

weakly developed crests, less massive zygomatic arches, narrower hard palate and some weaker teeth.

"Russian word

is

zarosshii;

lit.

"grown over"

Sci. Ed.

1342

Geographic

variation

of skull

insignificant,

manifested,

apparently, only in general dimensions.

Os
slightly

penis very large, quite massive and, on the whole, bent

upwards; anterior end forms slight bend towards ventral

side. Usually, also small

bend

in lateral direction.

broadens and

flattens, tapering anteriorly

At base, bone and becoming round in

cross section. At extreme end, there is a small thickening with a deep depression on lower surface. Body length of full adult (from four years old) and old individuals (Commander islands; 36), 100-136 cm (usually 120-130 cm); tail length, 30-36 cm; length of hind foot, 20-23 cm; length

of ear, 24-26

mm. One

especially large old kalan had a

body

length of 146 cm. Probably, in individual cases, body length

may

(Ognev, 1931; Barabash-Nikiforov, 1947, 1962; Stroganov, 1962). Females somewhat smaller than males (accurate
reach 150
data absent).

cm

females

Condylobasal length of male skull (31) is 124.2-145.9 mm, of (9), 120.2-142.9 mm; zygomatic width of males is 96.9116.2 mm, of females, 90.0-102.0 mm; interorbital width of males
42.0-46.5
is

is

mm,

of females, 36.7-42.2

mm;

postorbital width of

males

28.0-36.0

mm,

of females, 27.0-35.5

mm;
is

length of upper

toothrow (cheek teeth and canine) of males

females, 44.3-49.2

44.4-51.4

mm,

of

mm

(Ognev, 1931; Barabash-Nikiforov, 1947;

materials of Z[oological]

M[useum

of]

M[oscow]

U[niversity];

few specimens from Kamchatka). New-borns have body length of about 38 cm (Brandt, 1880) and weight of about 1.5 kg; "not long after birth" about 44 cm, tail length about 15.5 cm and weight about 2 kg; at two years old on average (36) body length 80.6 cm, tail length 29.7 cm, and weight 19 kg; at 3 years old, on average (75) body length 90.5 cm, tail length 31 cm and weight 21.2 kg; at 4-5 years old on average (17) body length 101.5 cm, tail length 31.6 cm and weight 23.8 kg. Later, dimensions and weight gradually and slowly grow and the fully adult animal often weighs from 23.0-36.4 (37) kg. Males may attain weight of 40 kg, while weight of adult females is 20-25 (30) kg. With age, relative tail length decreases, constituting 35% of body length in recently-born animals; in 4-5-yearold animals about 31%; and in 9-year-old and older about 25%
islands, a

Commander

(Barabash-Nikiforov, 1947, 1962).

Os penis length 150 mm, width

at

base 15

mm

(V.H.).

1343
891

Systematic Position

Only species

in the genus.

Geographic Distribution
Islands and in part shores of northern part of Pacific

Ocean from

approximately 62-64 N. lat. in the north, to Tropic of Cancer (in east [Pacific]) and 40 N. lat. on the south (in west [Pacific]).

891

Fig.

333.
line

Range of

the

sea

otter

or

kalan,

Solid

present

broken line reconstructed. Cross designates local settlement on Moneron Island. V.G. Heptner.
range;

Enhydra

lutris

L.

in

the

USSR.

1344

Geographic Range

in the Soviet

Union (reconstructed mainly ac-

cording to materials of Snou, 1902; Tikhenko, 1914; BarabashNikiforov, 1947; Uspenskii, 1955; Gribkov, 1963).

This occupied the southern coast of Sakhalin (apparently, Aniv

Bay and somewhat

to the north of

Cape Aniv along

Atlasov.

the

Okhotsk

coast)* and the whole Kuril chain from the southernmost islands

(Shikotan and Kunashir) to

In

Shumshu and

Kamchatka, along
in

its

western (Okhotsk) coast, sea otters

even

the

18th century, apparently, were present only in the

extreme south and did not extend north beyond the mouth of the Igdyg River (Ozernaya) and the Kambal' and Kitov capes (Steller,
1753, mentioned that in the "Penzhin River" sea otters were absent). In

any event, they did not

live in

Shelekhov Gulf, or perhaps

only very rarely intruded there (Steller' s data, given below, concern only the eastern shore of the peninsula). Frozen sea and fast

shore ice excluded their existence.

Along

the eastern shore of

Kamchatka

in the "Otter Sea", in

the first half of the 18th c[entury], kalans

tween 50 and 56 N.

lat.

were distributed bechiefly from Cape Lopatka to the Kronotsk

Peninsula (Steller, 1753), more precisely, to Cape Stolbov (a little north of the latitude of Bering Island about 56 N. lat.). Accord-

ing to less definite information, in the past (18th and first half of

19th c[entury]) they extended northward to 60 N.

lat. i.e.

approxi-

mately to Olyutor Bay and even to Cape Faddey and the mouth of the Anadyr' [river], in other words, approximately to 64 N. lat.

However, information concerning its occurrence at 60 N. and more to the north, is doubtful, or related to occasional
gressions of drifting.
892

lat.**

trans-

separate section of occurrence, not connected with the mainis

land portion of the range

ated at approximately 180

found

in the

Commander

islands, situ-

open sea from the nearest part of the Kamchatka coast. It is also isolated from the other part of the range which occupies the Aleutian islands about 350 km from
in the

km

Attn Island.

During the 18th, and especially the first half of the 19th was very intensive, entirely unregulated hunting of the animal throughout the whole range within, as well as outside,
c[entury], there

the boundaries of the

USSR. The

highly prized fur of the sea otter

Not

shown on map. Fig. 333 Contradictory Sci. Ed.

Sci. Ed.

1345
was, as

is

known, one of the main stimuli

later

for taking possession

of the northern coast of the Pacific Ocean, at

Russian explorers and

activities,

first, by individual by the "Russian-American Company".

In the second half of the 19th c[entury], after the

its

Company ceased
the animal

and

in

some
It

places

much

earlier,
its

was

practically exterminated in the greatest part of

region of occur-

rence in our country.

tal

completely disappeared from the continen-

coasts and Kamchatka, except

Cape Lopatka. On

the eastern

coast of Kamchatka, in Kronotsk Bay, the last time kalans were,

observed, apparently, was in 1852 (Ditmar, 1901).

At the beginning of our [20th] century (up

their

to

1943), otters

temporarily disappeared completely, apparently, from Lopatka, or

number decreased so

greatly that information of their exist-

ence there was questionable. However, in 1943, there were about

300 individuals (Averin, 1948); they are there now

time (beginning of the 60' s).

at the

present

On

the

Commander

islands, as a result of the senseless harvest

considered (Marakov, 1964) that kalans were completely exterminated and were absent here for about 100 until the 70' s of the 19th c[entury], when they were obyears served again. It is supposed that they resettled on the Commander islands across the sea. This is, however, quite doubtful. At least the present situation of populations of Mednyi and Bering islands points to the settled mode of life of the Commander kalans and to the low probability of their surmounting the large expanse of open ocean. On the first island, the otters are numerous and on the second they occur individually, although the islands are separated by a strait whose width is all of 24 km. Therefore, it is more probable that a very small number of animals was preserved from extinction on the islands and they, under conditions of complete prohibition of their pursuit, had the possibility of reproducing again. In any event, at the beginning of our [20th] century, there were already about 400 (Suvorov, 1912). Afterwards, they were mainly preserved because in this place, long before the Revolution, the state hunting of sea otters and fur seals was organized. On Mednyi Island, animals were always more numerous than on Bering Island. Otters had already disappeared from southern Sakhalin very long ago, and at the present time are absent there. On the Kuril islands, otters were from time immemorial, as is said, always present everywhere. However, pursued both by Russian and also many
in the 18th c[entury],
it is


1346
foreign fur companies in the
19th c[entury],
otters

quickly

decreased in number and their distribution was reduced. Their numbers continued to decrease in the 20th c[entury].

On

the brink for

two centuries, in part at the beginning of our century, they, although rare, occur on a quite considerable number of islands Shumshu, Paramushir, Onekotan, Kharumkotan, Ekarma, Matsuva, Raseva, Ketap, Chirpo and Kunashir (Tikhenev, 1914; Uspenskii,
1955).

On

the other hand, in 1912,

some authors believed

that otters

on the Kuril islands disappeared almost completely (Suvorov, 1912). Information concerning the 30' s (Barabash-Nikiforov, 1947), actually points to the

complete absence of

otters in the archipelago,

except for a few in Shumshu and between Onekotan and Urup.

Difficulty of access to
cially the small islands of its
least in very small

many

parts of the archipelago, espepart, permitted the animals, at

middle

numbers, to protect themselves

first

until the 40' s of

our century. In the very

893

post-war years

it

was established

that

sea otters inhabited Paramushir, Onekotan, Shiashkotan, the Musiru,

and probably several other small islands of the middle part of the
chain (Solov'ev, 1945; Sergeev, 1947; Kuznetsov, 1949).

By

the middle of the 50' s and the beginning of the 60' s,

i.e.,

within 10-15 years of initiating protection, the

in the Kuril islands

number of kalans
broadened; the

grew strongly and

their range

cause of

this

may be

only partially correlated, owing to better

censusing and inspection. In 1955, the population of Kuril kalans

was

in

any event not

less than

1500 individuals, and they lived on

Paramushir, Antsiferov Island (Shirink), Onekotan, Shiashkotan,

Rasshua, Simushir and Urup. South of Urup, animals were not

recorded (Klumov, 1957). After 1955, growth of the Kuril sea

and the expansion of their range continued, and in whole archipelago, were more comtowards mon south Urup (B.C. Voronov), and especially common on Urup. At the southern extremity of Kamchatka, sea otters were distributed (information from the 40' s and 1960) not only at Cape Lopatka; along the Okhotsk side, they lived to capes Kambal' and Kitov, along the ocean north to at least Gavryushin Kamen' Island (Cape Il'ya, Cape Zheltyi, Cape Inkanyushin, Utashud Island, Gavryushin Kamen' Island; Barabash-Nikiforov, 1947; Gribkov,
otter population

the mid-60' s, they occupied the

1963). All these places are located in southernmost Kamchatka, south of 52 N. lat. and are associated with Cape Lopatka. Kalans

1347

have now spread, however, and are considerably more to the north. Thus, in 1960, one animal was found in the delta of the Avacha river in Avachin Bay, and another (in winter) at the former Ust' Kamchatka region at 57 N. lat. i.e. north of this place, which
Steller (1753) referred to as the [range] limit.

The

latter

animal

was caught
1963).

in the river

20

km

distance from the sea shore (Gribkov,

Island, lying approximately

At the end of the 50' s, kalans were introduced to Moneron 50 km west of the southwestern end of
first

Sakhalin. The animals at

survived, but later died (G.V.

Voronov).
Therefore, the present range of the kalan occupies the

Com-

mander

islands, the

extreme southern end of Kamchatka (Cape

Lopatka and adjacent places) and all the Kuril islands. The rough number of animals apparently exceeds 4000 individuals.

Geographic Range outside the Soviet Union

Along the Asiatic coast, kalans were encountered on the coast of Hokkaido (Yeso), apparently, northern, eastern and southern and at the northern end of Honshu (Hondo, Nippon). These may in part be wandering animals (Temminck, 1847; reference to northern shore
of islands).
In the eastern part of the Pacific

Ocean, sea otters were present

along the entire Aleutian Island chain, in the Pribilof islands and along the American coast and adjacent islands from approximately 60 N. lat. (almost from the mouth of the Yukon at least along

the shore of the Alaska Peninsula) southwards to the middle part

of the California Peninsula [Baja California] (Sebastian Vizcaino Bay at Cedros Island, about the latitude 28 N. lat.; Hall and Kelson,

1959) and, perhaps, even Guadalupe Island. There

is

information

on
i.e.

their occurrence to the southern extremity of [Baja] California,

to the

Tropic of Cancer.

In the beginning of our century, in all

range, kalans were destroyed almost everywhere and were

American sections of the met

with in very small numbers only along the Aleutian chain and

perhaps individually

in

the San Francisco region* and in other

separate places. In San Francisco, where sea otters were consid-

ered to already have been extirpated long ago, they unexpectedly

Actually, Monterrey and Big Sur, south of San Francisco Bay

Sci. Ed.

1348

894 Fig. 334. Reconstructed range of the sea otter, or kalan, Enhydra the coast of North America. V.G. Heptner.

lutris L.

on

appeared
in

in the 30's in quite considerable

numbers (Bolin, 1938;

Fisher, 1939).

Thanks to their protection, the population of otters America quickly grew; at the beginning of the 60's they were

already quite significant. Consequently, their range has also wid-

ened. The question

now concerns

organizing utilization of the

animals (V.H.).

Geographic Variation
Geographic variation
materials of skins in
in the

kalan

is

poorly understood. Collection

museums

are very

few because of

their very

high price, and these skins are often old and

894

their color has

changed.

Conceptions of species variability

are, therefore,

based mainly on

craniological features and only partially on color and fur characters.


1349

Within the USSR, two races may be

1.

distinguished"*.
I.

Commander

sea otter, or kalan. E.

lutris

Linnaeus, 1758

(syn. marina).

Fur color relatively

light,

with significant development of gray

hair and quite long guard hairs (30

long on the average, in middle of back). Skull fairly large, moderately wide. Zygomatic width constitutes 71-80% and mastoid, 71-79% of condylobasal
length.

mm

Condylobasal length of skull of males (17) is 130. 5-M 136.5140.3 mm, of females (7), 120.0-M125.4-128.2 mm; zygomatic width of males is 101. 0-M 104.3-110.0 mm, of females, 90.0-M
94.3-102.0

mm;

mm,

of females,

interorbital width of males is 40.2-M 43.4-46.5 36.7-M 39.2-42.2 mm; mastoid width of males is

93.9-M 99.5-108.0 mm, of females, 88.0, M 91.4-95.3 mm; length of upper toothrow in males is 46.2-M 48.3-51.4 mm, of females,
44.3,
895

46.0-48.6

mm.
is

Greatest skull length

Nikiforov, 1947).

144.2

mm

on average (Barabash-

Commander
Outside the

Islands.

USSR, Aleutian and

Pribilof islands, and on the

American coast, from the northernmost place of occurrence south to Vancouver Island inclusive.
2.

Kuril otter, or kalan. E.

I.

gracilis Bechstein,

1799 (syn.

crientalis, stelleri, kamtschatica).

relatively short guard hairs (on

Fur relatively dark, with weakly-developed gray hair and a middle of back, about 25 long

mm

on average).
Skull

what

flattened.

somewhat smaller, wider than in nominal form and someZygomatic width constitutes 85-87%, mastoid

84-86%

of condylobasal length.

132. 8-M 133.4-133.6

Condylobasal length of skull (5 old males; Kamchatka) is mm; zygomatic width is 113.3-M 114.9-

116.4

mm;

interorbital width is

46.9-M 48.0-49.1 mm; mastoid

however, slightly

"Characteristics of the races after Barabash-Nikiforov, 1947 and Stroganov, 1962.

Description of fur of different races by Barabash-Nikiforov, 1947

is,

contradictory. In contrast to other authors, he indicates (p. 26) a "tawny sheen" on fur

of Kamchatka animals, giving fur even a "reddish" color, compared to fur of

Com-

mander animals. On the other hand, he refers to a "darker tawny tint" of fur of Kamchatka animals. The review of Stroganov (1962) does not give anything new and
only complicates the picture
the species).
(it

refers to the existence of "not less than five" races of

From

this

work,

if

not with reservations, data on dimensions and propor-

tions are taken.

1350
width
is
is 107.0-M 109.5-112.3 mm; 46.2-M 46.6-47.0 mm.

the length of upper toothrow

Greatest length of skull

is

137.3

mm

on average (Barabash-

Nikiforov, 1947).
Kuril islands, Kamchatka, and in the past, in South Sakhalin.

Outside the

USSR in

the past, northern Japan (Hokkaido,

and possibly northern Honshu).

and Kamchatka sea otters into separate races (Stroganov, 1962), must not be considered justified. Those geographical considerations on which this was based do not give any basis for this. There is no isolation between kalans inhab-

The attempt

to separate Kuril

iting different parts of the chain;

nor

is

the

Kamchatka population

isolated

from the Kuril. Sea otters not only drift with currents and storms (to which even Steller, 1753 referred), but they themselves undertake quite large migrations and undoubtedly move from one island to another. Apparently, the populations of Cape Lopatka and

communicate with each other^ In particular, this from information given above concerning the "disappearance" and "restoration" of the Lopatka population during the

Shumshu
is

Island

inferred

last ten years. It is also

known

that in the

pre-war years, Japanese

kalan hunters tried

more than once

to "drive"

Lopatka animals

to

Shumshu using various means. Kuril kalans were once considered the very best and were especially valuable. Kamchatka were also esteemed more highly than

Commander
and

[otters].

This correlation of values corresponds to

ra-

cial difference

between Kuril-Kamchatka and Commander

[otters]

to a certain degree, indicates the isolation of the

Commander

population.

Outside our country, only one well distinguished form is apthe North American parently known: E. I. nereis Merriam, 1904

coast south of Vancouver Island (V.H.).

Biology
Population. In the second half of the 18th century, populations of
kalans were
still

significant in

some

places.

Concerning

this

one

may judge from

the fact that on the Pribilof islands, in 1787,

more

than 5000 kalans were caught. Later, their number greatly decreased,
'Width of the
passage
1947).
is

First

Kuril Strait, separating

them "reaches 10

miles, but free

reduced to 3.0-3.5 miles due

to reefs present

on both shores" (Sergeev,

1351

from

killing

and

in

1821 the harvest was prohibited, and by the

896 40's they

completely disappeared here. During 1763-1764 on the Andreyanov islands, about 3000 kalans were taken, and during

1775-1780, on Urup (Kuril islands) 1170. After the strong earthquake of 1780 kalans disappeared here. With renewal of harvesting on the Kuril islands, during 1828-1830, 2600 animals were caught on only Urup and Simushir. For 1842-1860 on the Kuril islands, 4510 individuals were taken overall, i.e. an average of only 250 animals per year. In 1745, on Bering Island more than 1500 kalans were captured, and from 1747-1749, an additional 1500. As a result, for the period 1754-1755, a total of 5 kalans were caught, and in 1756 not one. In 1762, 20 head were taken

and

after that, the harvest

was suspended

for a long time in

view

of the trifling results

kalans were very few. Therefore, on Bering

Island during the second half of the 18th century, the stock of

kalans was slightly more than 3000 head. With depletion of the

on Bering Island, the main harvest was shifted to Mednyi Island. Here, in 1754, no less than 790 otters were caught, and during 1760-1763, no more than 20. Here, probably, the population was half that on Bering Island. Overall, apparently, only 40005000 of these animals inhabited the Commander islands, since there is information that the quantity of kalans on Mednyi Island
otter stock

attained 2000.
for

From

these uncoordinated figures (complete data

that at

whole range are absent), the conclusion may be drawn,

the beginning of the 18th century, the kalan population consisted

of approximately 20-30 thousand (possibly slightly more) in the

whole range

from the

northern coasts of Japan to the shores of

[Baja] California. In the latter area, during the past century,

7000

skins were bought from the native hunters in only one year.
In order not to undermine the basic stock, not more than 20003000 animals could be harvested every year throughout the whole

range. In the period beginning in 1870, the world take of kalans

4000 to 8000 head, on average 5000 head per year. Therefore, the entire time harvest was always at the partial expense of the basic stock, and the result was that after the
fluctuated within the range of

harvest of 8000 head in 1884,

it

steadily decreased; in 1900,

it

was

1000 head. This decrease continued until the nearly complete liquidation of legal harvest in the beginning of this century. Within more than two centuries, otters were almost destroyed due to their rapacious exploitation.
to

reduced

1352

On

the

Commander

islands, after the period of almost

com-

plete disappearance of kalans, they appeared again in the 70' s of

the previous century, but only on

Mednyi. The main stock of Mednyi

in

Island settled mostly on the northern (northwestern) extremity of

the island. Disturbed

by a take of living animals

again
to

1937-1938, the
1950).

herd migrated to the southeastern extremity

later,

(Il'ina,

Ten years
of

kalans

returned

the

northern

extremity

the island. If in 1931-1932, 282 kalans were counted in the waters

of

Mednyi

Island, then in

1952, 250 animals were recorded

(O. Danilov), and in 1955 about 350 (Marakov, 1964). Therefore,

despite the absence of harvesting due to

difficult to eliminate, the

some reason which were

Commander

stock of kalans hardly grew

during the
In

last

20 years.

1924, on both

counted; in

1939800; and

Commander Islands, 400 individuals were in 1957500 (Nikolaev, 1958). Dur-

ing the last decades, the situation sharply changed, and the kalan

population began to grow quickly

in

1964, on Mednyi, the number

of adult animals alone reached 1000 (Marakov, 1964).

As shown

above, the population of sea otters on Bering

is

very small; they

'^

m^^z

^^^^.
897
Fig. 335. Habitat of kalan

.-^

'^^^^"-^^Mife^
'

on Mednyi Island. Photograph by S.V. Marakov.


1353
are encountered individually.

It

follows that growth in the number

of animals
ity

is

correlated to their reproduction locally

is,

the

possibil-

of their immigration from outside

It

in practice,

very improb-

able (Marakov, 1964).

must be noted

that recent

methods of
state fur

counting

still

allow significant discrepancies between data of vari-

ous authors. Thus, in 1931, according the

Commander

farm put the number

(Barabash-Nikiforov,

at

282, but from another source of data

500.

1948) about

Kalans of the Kuril chain and Kamchatka constitute one population,

which

is

characterized by regular seasonal migrations

(Dobrovol'skii, 1936; Nikolaev, 1958). In the Kuril chain, there

were 200 in 1912; in 1924600, and in 1939800 kalans. By autumn of 1958, north of Iturup there occurred about 1900-2200 animals living; and by 1961, the population increased to 29003100 head. On the islands of the archipelago inhabited by kalans, they were most numerous on Urup 938 animals in 1958, and 1700 in 1961, and on Paramushir where in 1958, 614 individuals were recorded. In the remaining cases, the number of recorded animals did not exceed 79 (1958). Relatively more animals were observed in the waters of Shikotan, Ketoy and Simushir islands (Nikolaev, 1958, 1964). On Kamchatka in 1910, about 306 sea otters were censused, in 1924 400, and in 1943 again only 300 (Averin, 1948). The total number of our kalans at the beginning of

the 60' s, apparently, exceeded 4000.

On

the islands of the Aleutian chain and the shores of the

Alaska Peninsula

in 1912, there were about 200 otters; in 1924 200-250; in 1958, thanks to rapid growth of the population, the number had already reached several thousand. In the waters of the

California coast (36 parallel and south), in 1938 in

all

94 were

censused; but
the are

in

1958, already there were about 500. According to

most recent data, along the coast of the state of Alaska there 20-30 thousand otters. It is calculated that on the islands of the
its

Aleutian chain, in particular on the Rat, islands, the kalan population has already reached

density limit (Kenyon,

1961). In

1962, with examination of six main sections of occurrence from

11000 kalans were counted (Brooks, 1963). Attempts to animals on the Pribilof island have still not succeeded. Proceeding from the given data on America and paying attention to information on the animal's population in the USSR and in California, the total number of kalans may be determined as
the
air,

settle these

1354

Fig.

336.

Biotope of kalan.

Babichii

Pod'em Bay on Mednyi

Island.

Photo-

graph by S.V. Marakov.

25-35 thousand head. Apparently,

otter
is

this species' extinction, anticiit.

pated not long ago, no longer threatens

Protection of the sea

one of most remarkable

in the field of nature protection.

Habitat. Habitat of the kalan

898

shores, barrier reefs,

is characterized by steep rocky submerged and exposed rocks, continually

washed over by waves,

as well

(species of Alaria and Laminaria

"sea cabbage") extending

The
latter

as

vast thickets of sea

weed
as an

interrupted belt along the shore.

mer, but are destroyed by storms in winter.

grow strongly in sumEverywhere where they

exist at the present time, kalans prefer the extremities of islands

and peninsulas. Here the animals have the possibility of moving around to the lee side in stormy weather. Condition of the water medium, feeding and climate are in
total

more important

for kalans than the character of the coastal

belt itself. Their inclination

toward inaccessible coastal zones de-

1355

veloped as a result of savage pursuit. In structure and type of

adaptation, the animal
to living
is

not only littoral but also pelagic, adapted

under conditions of the northern part of the Pacific Ocean.

and cool summer, and and entirely variable, but occasionally severe, winds especially in winter. These winds, and especially storms, very unfavorable for kalans. The small difference are between air temperature and that of the upper layer of water seems to be a positive factor, especially in summer. The difference in
This
is

a marine climate, with mild winter

fog, frequent drizzle

water temperature during a year on the

Commander

islands

is

only

15C
to

in all

(from 1.5C

in

January to 13C in August). In the

Kuril chain, water temperature fluctuates from

1.8C

in

January

5-1 0C in the northern islands and 10-20C in the southern

islands.

The

lag of the period of greatest coolness at a depth of

20

and more, has large significant in the ecology of marine animals

serving as food for kalans. During experiments on acclimatization

on the Murmansk coast (Reshotkin and Shidlovskaya, 1947), kalans suffered some from summer heat and mosquitoes.
Food. On Mednyi island, two species of sea urchin serve as the main food of kalans; bivalve mollusks are of significant importance

octopuses are
When

less so. In

constitute

10%, but fishes

6.7%,

numbers, crabs of several species of which cod, lumpfish

{Cyclopterus lumpus]. Pacific capelin, sand eels [Ammodytes] and

sockey salmon predominate. Altogether, 28 food components were

established.

held in captivity, the kalan reveals great frugalnatural conditions,

its

ity in feeding.

Under

food regime

is fairly

monotonous (Barabash-Nikiforov, 1947). The chief characteristics of the animal's nutrition in tumn-winter period on the Commander islands consist of
899 that at this

the authe fact

time there

is

a sharp fall in the proportion of crabs

to

(from 24 to 4%) and fish (from 22

6%) which migrate

at that

time from the shores into great depths of water. The proportion of

mollusks slightly decreases, from 50 to 45%. Sea urchins remain

the

main food component (100-98%). On the Pribilof

islands, sea

urchins also constitute

is

92%

of the kalan's food (Murie, 1940). This

otters in nature,

also confirmed

by observations on

on Mednyi

Island (Barabash-Nikiforov, 1947). On Urup February-March, kalans in the western (Okhotsk) coast feed on sea urchins, polyplacophore, gastropod and cephalopod mollusks, crustaceans, starfish, fish and algae. Along the eastern coast at this
(Kuril chain), in

1356

Fig. 337. Zapalata

Bay on Mednyi Island

place of residence of kalans. Photograph by S.V. Marakov.

time, crustaceans

occupy

first

place, various mollusks

and then

sea

second,

urchins and algae (Nikolaev, 1958).

In captivity, the daily [dietary]

norm of

the kalan constitutes

4.5-5.5 thousand calories (Reshotkin and Shidlovskaya, 1947).

According to other data, for the average sea otter, it is 6.5-7.5 thousand calories, but for large animals even 10 thousand (Il'ina, 1950). Proceeding from the fact that the quantity of sea urchins eaten by a kalan in one day contains only a total 3 thousand large calories, some (Il'ina, 1950) consider that, in freedom, fish must predominate in the kalan's food, and kalans cannot feed on sea

urchins alone. In captivity, food of the kalan consisted of fish

(66.8%) and sea urchins (33.1%). As concerns

fish proportion increases to
(Il'ina,

digestibility, the
to

80.3%, and sea urchins drops

19.6%

1950).

It

is

impossible to consider that the question of

kalan nutrition has been sufficiently clarified.

Home

range. Kalans do not reveal an attachment to any sort of

definite place. Individual

animals and groups of them appear

irregularly in first one place and then another; however, true

1357

Fig. 338.

One of

the groups of kalan of

Mednyi

Island.

Photograph by S.V. Marakov.

migrations are only observed in separate individuals and pairs.

Periodic redistribution of the kalan populations in the region of

900

permanent habitat which happens locally, depends first of all on the weather. With a wind of velocity 4, several concentrations of them are observed at the shore. This becomes very obvious with wind velocity 7. In calm summer weather, separate kalan groups stay permanently at a distance of 18 and more kilometers from shore, where there are shallow places (banks), with beds of sea cabbage [kelp]. Here they spend the night. In winter, kalans gather themselves at the shores, since the kelp beds are destroyed by storms. In the open sea, kalans may be met with at a distance of 18-27 km from shore, but they are capable of reaching considerably greater distances. The appearance of killer whales* (which usually cause otters to flee in panic), human activity and the extheir

haustion of main foods, are reasons for local regrouping of kalans;

such was the case in 1938, when on Mednyi Island sea urchins

almost completely died out following some sort of epizootic (Barabash-Nikiforov, 1947).
*Russian name
nal
is

"kosatka", misspelled "kasatka", or swallow, in Russian origi-

Sci. Ed.

1358

Among

kalans of the Kuril chain are distinguished local ones

in

which are permanently present

migrating ones.
In

one and the same place as

summer, kalans here completely confine themselves in defisometimes move for tens of kilometers in search of food and protection from storms. Burrows and shelters. For resting, kalans utilize sections of rocky laida, narrow coastal strips under steep coastal cliffs, especially in times of winter storms. For the same purpose, they very frequently use the emergent and submerged rocks ("hidingplaces*") at various distances from the shore. In summer, otters also frequently use dense kelp beds, which are sometimes considerably far away from the shore, for resting and protection from the
nite sections of the shore, although they

attack of killer whales.

Daily activity and behavior. The kalan

active during the day.
it,

is

an animal preferably
or shortly before
active only on rare

Its

activity begins at

dawn
is

and ends

at twilight.

At night, the kalan

occasions, usually on moonlit nights. In winter time, the animals

901

spend the night on shore and eat the entire day without an interval

Fig. 339.

Kalan lying
is

in water,

May
Ed.

1960. Photograph by S.V. Marakov.

*Russian word

tainiki

Sci.

1359
in the activity for resting. In summer, they feed mainly in morning and evening, resting from 11-12 to 15-16 o'clock. In this time, the animals stay in the algae beds, gathering in herds of 20-120 head. Old otters, sometimes "twist", wrapping themselves in algae and appearing as if anchored (Fisher, 1939). The degree of activity of the kalan depends upon the provisioning of the given region with food, the weather conditions

and the

state of the sex glands.

Animals are more active

in regions

with small food supply. Males also manifest increased activity in

search of females in estrus. Before storms, activity of kalans also

grows
itself,

in

connection with relocation to protected places. The kalan

it

feeds several times in the course of a day. After satiation,

cleans

sometimes plays, and then

rests in the water, lying

on

its

back. In the open sea near shore, the kalan

may

feed even with

wind strength of 6-7.

In the sea, the sea otter In searching for food,
it

is

agile,

mobile and quite quick.

moves at a speed of 5-6 km/h, but its maximum speed equals 12-16 km/h (A.M. Nikolaev). Capture of

902

Fig.

340.

group of resting otters on the rocks of Glinka Bay, Mednyi Island, December 1958. Photograph by S.V. Marakov.


1360

bottom and swimming fish is with its teeth and fore paws, then clasping them to its chest. Coming to the surface, the animal lies on its back with it and in such a position eats it. This pose is very characteristic for the animal in feeding and resting. After taking food, the kalan cleans itself of the remains by revolving around on its [body] axis. After resting, kalans often gambol, play with each other and somersault in the water. The kalan is a herd animal, and peaceful for the most part with respect to each other. However, fights between males for females occur in places. On land, kalans move quite slowly, strongly bending their back while walking, like the river otter. Droppings on land with abundant remains of sea urchin tests and remains of crabs, mollusks and fish serve as a
characteristic feature of the kalan's residence.

Seasonal migrations and transgressions. Until recent times,

it

was considered

do not perform significant and regular migrations. This idea was based on observations of herds of the
that kalans

Commander
902

islands. In actuality, a significant part of the Kuril

(perhaps Kuril-Kamchatka) population gradually translocate themselves from wintering places on Iturup and
to a place of spring residence

Urup

islands northward

on the middle islands of the chain

Brouton, Onekotan

where

the animals spend

March and
perhaps

April.

Then they move

islands

farther to

summer (May-October) on
Aland
and

the northern
to

Paramushir,
coast.

Shumigu,

the

Kamchatka

In autumn, a local reverse migration to the south takes place.

Autumn migration
is

of kalans from the northern islands of the chain

associated with decrease in water temperature toward -1.8C,

which causes displacement of the bottom fauna to deeper places, complicating food capture. Another cause leads us to think it is the appearance of ice which hinders movement and food-catching. At the end of January, cooling of water to -1.8C and the appearance of ice drive the kalans away from the shores of Iturup Island and in March, also from the Urup coast. The coastal waters of the middle islands remain free from ice in March-April. In the time of migration, kalans swim singly and in pairs at intervals of 10-15 min[utes]. During migration, all animals securely hold to a definite direction, neither stopping nor paying
attention to local kalans. Nevertheless cases occur of single indi-

viduals (possibly young)

who

lose their

way and

return (Nikolaev,

1958).

1361

903

which do not include the there was observed translocation of a herd for a considerable distance, after which the animals reappeared in the abandoned places, sometimes only after a series of years. Such was the case when kalans disappeared from Urup after the earthquake in 1780, and also a series of cases when the disappearance of kalans in one region was accompanied by sudden growth in the population of another. In the 40' s of the previous century, otters disappeared from the shores of Simushir Island after the eruption of Prevo volcano. At the same time, they appeared on Shumshu, Rasshua and Ushishir islands, where they absent earlier. Kalans temporarily left Cape Lopatka after the wreck
In addition to seasonal migrations
entire kalan population,

more than once

Fig. 341.

Females with young and pregnant female otters. Mednyi Island, Zapalata Bay. 6 August 1962. Photograph by S.V. Marakov.

1362
it is sometimes believed possible (Barabash-Nikiforov, 1947), to connect the secondary appearance of sea otters in the 70' s of the last century on the Commander

of a Japanese ship. Finally,

islands (after their complete or almost complete destruction there by the end of the 18th century) with increase of the harvest at that time on the Aleutian and Kuril islands, which might have caused their resettling on the Commander islands. As shown above, this

idea

is

lacking in foundation. There

is

the suggestion that all kalan

populations, with the possible exception of the isolated California

group, constitute one chain, within the boundaries of which displacements and regrouping can, apparently, occur.
the third year of

Reproduction. The kalan, apparently, attains sexual maturity in life, but this is not accurately established. The
at

kalan has no definite estrus (rut) period. Nuptial games, mating

and new-born young with the females are observed

the year.

any time of

was somewhat more often observed in spring March-May, but it was also observed in JulyAugust. On the Kuril islands, the mass parturition proceeds in

On Mednyi

Island, nuptial play

spring, but

new-borns are also observed

at

other times of year

(Nikolaev, 1958).

Fig. 342.

Female kalan with

ihe

new born on

its

chest.

Mednyi

Island.

Photograph by

S.V. Marakov.

1363

904

The duration of pregnancy is determined as 8-9 months; in one case, it lasted 238 days. The female, as a rule, bears one cub. two cubs In very rare cases, two embryos were found; still rare

with a female.

tion

3-9 min[utes] water and Mating takes place on the shore or on rocks flooded with water. Delivery
in
lasts

parturilasts

about two hours. In several cases, the cub of the previous litter, already almost equal to her in dimensions, is still with the parturating
female. Clearly-defined [mated] pairs are absent

among

otters.

The annual increase of the kalan population on Mednyi Island was determined in the 20's-30's of the present century as 7% or less (Barabash-Nikiforov, 1947). These figures characterize the tempo of growth of a herd strongly undermined by predatory harvest.

Rate of annual growth

may be judged by

the percent of

yearlings in the population. According to data of the

autumn count

growth was 10.5%, Onekotan 6.6%, 8.5%, Shiashkotan 14.0%, on Ketoi and Simushir islands Urup 7.8%. Therefore, the probable annual growth of the Kuril
of 1958, on Paramushir, the

Fig.

343.

Otters

going

to

water.

Mednyi

Island.

May

1958.

Photograph by

S.V. Marakov.

1364
population fluctuates within limits of 6.6 to 14.0% (A.M. Nikolaev).

Average equaled 9.2%, which

figures given.
It is

is

somewhat higher than

limits of 10 to

earlier

undoubted, that in the thriving period of the spehigher

cies, natural increase is

within

20-25%.

Growth, development and molt. Immediately after birth, body length of new-born is 50-56 cm, and weight, 1.4-1.6 kg. Kalans are born able to see, with juvenile fur and having 26 milk teeth. In the first days after birth, they are barely active, and lie on their backs on the shore or in water. First attempts to swim independently are

made at the age of about 2 1/2 weeks, and by the third week, they are already able to swim for short distances. The juvenile (infant) fur is replaced by permanent at the age of about 6-7 months.

905

At this time, replacement of milk incisors and first premolars by permanent is completed. Replacement of second and third premolars begins, and the molar teeth erupt. Total number of teeth is already 32. Body weight of such a "koshlak" reaches 9-1 1 kg, body length, 1.10-1.15 m. At one year, live weight reaches 17-19 kg and body length, 1.25-1.35 m. Replacement of premolar teeth is completed, but skull crests are still weakly developed (Il'ina, 1950). Enemies, diseases, parasites, mortality, competitors, and Dynamics of number. Among animals, the most dangerous and almost the sole enemy of kalans is the killer whale which often manifests

The polar shark is rare and is mainly found Old information, that Steller's sea-lions and fur seals also appear to be enemies of the sea otter, have not been confirmed by more recent observations. The largha seal is a competitor of the kalan as regards main food components, and in respect to shelters, Steller's sea-lions, fur-seals and other pinnipeds
itself in otter habitats.
at great depths.

are competitors. Several of the sea birds are

tors, in

shown

to

be competi-

small degree of the kalan. However, their activity has no

fundamental effect on food resources of the kalan. Natural mortality of kalans is very great. Causes show up as age limits, diseases,
injuries

caused by

killer

to shore laidas; the breaking of ice (Nikolaev, 1958),

whales and, possibly, sharks; rockfalls on and in indi-

vidual cases, the effect of storms and birth pathologies.

Age

limit

of the kalan

is

not known. Judging by the relationship between the

and duration of life, must be considerable. There are no reliable data concerning population dynamics of otters. For the last 200 years, the human role was so great that other factors were overwhelmed and could not be estimated.
in kalans
it

intensity of reproduction, animal dimensions

1365

Besides direct destruction, the movement of ships in the region of their habitat and frequent frights caused by other
activities

human

have an extremely negative effect on the numbers and of

kalans and their colonies on the coastal belt.

906

Practical Significance

The economic significance of

the kalan

is

very substantial. This

durable fur and
is

now

rare animal provides beautiful,

warm and

considered the most valuable fur-bearing animal. Individual otter

London auction for a price of $2000. The value of the fur is doubled by the rarity of the animal: for the past 80-100 years, not more than 8000 hides were sold per year,
hides have been sold at the

and

in the past

30 years

a negligible quantity (single ones), and

more or

less accidentally.

bition continues until the present with the

was prohibited in 1924. This prohiaim of future growth and dispersal of herds. Because of the low reproductive rate of
In the

USSR,

the harvest

kalan and conditions not yet entirely favorable for them;

present time, conditions for restoration of
quire sufficiently
initial

at the

population re-

more

time. At the present time, the question has

been raised concerning a possible catch of a limited number of otters, especially the sick and old, for skins (in nature, mortality in this species is relatively very high: Marakov, 1964). Along the

American

coast,

where the density of

its limit,

otter colonies locally has

already reached

a test harvest

was conducted
in the

in

1962 and

174 animals were caught (Amchitka Island);

there

following year,

was planned

a harvest of

up

ing the possible price in the fur

300 individuals for determinmarket (Brooks, 1963).

to
in

Several perspectives

may be borne

mind concerning cage

in captivity

rearing of otters; experiments to keep

relatively successful.
ral

them

have been
their natu-

However, feeding the animals on

food

in captivity is very
is

complicated, since preparation of this

food which

needed in large quantities for each animal is difficult. The experiment of the keeping animals in open-air confinement on the Murmansk coast of the Barents Sea, may be also
considered partially successful.

The main means of catching otters in the shutter net, and in summer, the landing-net. At the epoch of abundance of this animal, harvesters caught it on land by hitting it on the head with a
stick (P.Yu.).

907

Supplement

to the

Order Carnivora

Superfamily of Wolves
Superfamilia Canoidea Simpson, 1931

Family of Raccoons
Familia Procyonidae Bonaparte, 1850
Predators of small, moderate and large dimensions, in part generalized,

and

in part specialized

and highly specialized types'.

Limbs
ity

are pentadactyl, plantigrade and semi-plantigrade; digits

separated to different degrees, the third

longest; claws
some

in major-

of cases non-retractile, in a few, semi-retractile (Ailurus, panda;

Bassariscus, cacomistle). The lower surface of foot and hand usually

bare.

General form of skull variable

skull strongly elongated.

in

species facial region

still

elongated, in others, skull short and rounded; in

others,

whole

However, in the majority, braincase relasometimes very, voluminous. In majority of species, alisphenoid canal is not developed, canal of carotid artery (can. caroticus) located at inner side of auditory bulla and separated from posterior lacerate foramen (for. lacerum posterius).
tively,

Dental formula in majority

is

I-C-P M = 40,

14

in

some,

I-C-P-M- = 38.

3132

(Ailurus, red panda) or

I-C-P-M- = 36
I-C-P M = 40
3

3132
14
2

(Potos, kinkajou or monkey-tailed "bear"), or

(Ailuropoda,

bamboo

bears or giant panda).*

'Characteristics of the family, and later of the genus, are given briefly.

*Now

recognized as a true bear, not a procyonid

Sci. Ed.

1367

Form

of cheek teeth and character of their masticatory sur-

faces extremely variable. In one, they are powerful and broad with
a flat multicusped, bluntly tuberculate masticatory surfaces, typical of

omnivorous and phytophagous carnivores (extreme form

is

Ailuropoda, bamboo bear). Teeth in several are characteristic of

meat-eating carnivores, of carnassial form with cutting crests. The
fourth upper premolar and the first lower molar are transformed
into carnassial teeth, resembling teeth of species of wolf family, Canidae (Bassariscus, cacomistles). In majority of species, cheek relatively broad and flat. Incisors rather teeth of intermediate type weakly developed, the first premolar has tendency toward reduction small or absent, sometimes disappearing with age. Canine

908

relatively small.

The OS penis

is

present and well-developed. Anal region de-

void of glands and glandular areas.

are quite varied

As regards general appearance and size, species of with a small number of species in

the family the group,

differences are considerably larger than in other families of the

order.

One

of the general aspects and dimensions

is

bear-like

{Ailuropoda,
panda), others

bamboo

bear),

some

are similar to cats (Ailurus,

fox and raccoon-dog (Procyon, raccoons) or weacacomistles) and

sel (Bassariscus,

coati; Potos, kinkajou or

some are quite monkey "bears").

is
still

individual (Nasua,

In

some

species,

muzzle

short and obtuse; in others,

it is

of

the fox or Arctic fox type, in

others, with very long pointed

facial portion; in the majority of cases, ears quite large, rarely

short and rounded; eyes usually quite large. In majority of species,

tail

long; in some, longer than trunk, covered with dense luxuriant

hair; in

one species, very short

(as in bears), in another, strong,

tail

muscular and prehensile like a monkey's

rare exception

{Potos, kinkajou; a

among

carnivores). Pelage quite long and dense; in

some, very dense, fluffy and long (raccoon, providing valuable fur). Color entirely variable from more or less monotone yellow or brownish to very mottled and bright, consisting of a combina-

tion of bright red with black {Ailurus, panda). In a series of species,

dark transverse rings are well set off against the light general
tail.

tone of the long, fluffy

Differences in dimensions very great

length of about

smallest forms have

in

body

25-30

cm

and about 1.25 kg

weight {Bassariscus,

1368
cacomistles), and in the largest, head and trunk length

is

about 180
bear).

cm

and the weight

is

about 150-160 kg (Ailuropoda,

bamboo
with

Species of the family are mainly in tropical,

in part

warm
winter

temperate climates, and only one (raccoon) extends quite far into temperate zone, and
sleep.
in the north

of

its

range even

falls into

Some

species ascend high into mountains and are even

com-

pletely associated with

montane regions (Asia

see

below). All

are, to a great or less extent,

forest animals or associated with

forest regions, the majority closely associated with forest, climb

trees well

and often, some live

in tree

hollows, etc.

series of

species are mainly, arboreal animals (semi-retractile claws) or even

specialized in climbing (monkey-bear, Potos).

Some

species are

very fast and active, others are languid and slow in their move-

ment. The majority are solitary nocturnal animals, forming only

temporary family groups. One species is partly diurnal and sometimes forms large groups (Nasua, coati). The majority are omnivorous animals, chiefly "gatherers", feeding in addition to plant foods, on invertebrates and lower vertebrates, small rodents, birds, etc. Some are more predatory, a few are mainly phytophagous or specialize strictly on plants (bamboo bear, feeding on young shoots of bamboo). The range of the family is divided into two parts a vast American and relatively very small Asiatic. In America, the range occupies a considerable part of North America, except its northern part; Middle America; and a large part of South America, except its southern part. The northern borders of the range extend from the Gulf of Saint Lawrence, through southern Quebec, southern Ontario, the south of Manitoba and Saskatchewan, Alberta and through the south of British Columbia (for details, see below; description of raccoon). The southern border runs from the Pacific

910 coast of northern Chile (at approximately 20 S. lat.) to southeast

to the

mouth of

the

La

Plata [river] on the Atlantic coast.

The

range includes Vancouver Island and several small islands lying

directly along the shores of the southern part of North

America,

some of

the

Bahamas and Lesser

Antilles islands (see below, spe-

cies description)

Himalayas
region).

in

and Trinidad. In Asia, the range occupies the Nepal and Sikkim, and farther to the east upper Burma,
north to the border of

Yunnan and

Gansu (an

entirely

montane

1370

910

Fig. 345.

Range of

the raccoon genus,

Procyon Storr*. V.G. Heptner.

Sci. Ed.

Misspelled "Starr"

in

Russian original

1371

The family Procyonidae

ted into
it,

in all of the diversity of

forms admitis

according to the present point of view, represents a

suffi-

systematically diagnosed and well-defined group, which

ciently connected

by

common

origin. Actually, this

is,

in substance,

the broadly accepted opinion.

The

diversity of forms within the

family must, apparently, be considered as an adaptive divergence, associated with feeding characteristics, and different degrees and

forms of adaptations to climbing.

911

At the same time, diversity of structures within the family, led and sometimes still leads, to strong fragmentation. Separate authors divide it into 2, 3 and even 7 separate families. In the latter case, each well-defined genus is separated into a distinct family. There has also been no lack of division of the family into a series
of subfamilies.

For example, several authors not only separate the genera Potos
(kinkajou) and Bassariscus (cacomistle) from the family, putting

them

in the

rank of separate families, but also entirely rejected the

idea of close relationship with remaining Procyonidae.

tematic position of the genus Ailuropoda

The

sys-

and

is,

frequently treated in various ways.

(bamboo bears) has been, It was related to bears,

as a

separated into a special family (alone or together with the panda,

Ailurus), and only relatively recently,
it

was again recognized

member

of the raccoon family.*

far, the

Having gone

subdivision of the raccoon group, as with

utility
its

anything to such detail, does not bring

and only obscures

relation to others.

the general picture of family structure and

Evidently,

more natural to consider a single family, Procyonidae, composed of two recent subfamilies (see below).
it

is

In the circle of families of the order,

and the group Canoidea,

the family Procyonidae

shows the
is

clearest relationship to the bear

family, Ursidae (see characteristics of this family). Relationship to

the wolf family, Canidae,

regarded as

much

less close,

although

some data

also point to a certain relationship

between these groups

(see characteristics of wolf family, Canidae, and of genus of rac-

coon dogs, Nyctereutes). Also, there appear to be no clear relationships between the raccoon family and the marten family, Mustelidae.
In the series of families of the order, the raccoon family
is

usually

"Presently

Ailuropoda

is

considered a bear

Sci. Ed.


1372
placed as the immediate neighbor of bears and between bears,
Ursidae, and martens, Mustelidae.*

The raccoon family

and
is

is

less ancient than the

its

wolves, Canidae,

associated, apparently, by

origin, with early

The main American branch of the family split off in the lower Miocene in the Americas, and throughout its entire history, it was restricted, as at present, to South and North America, generally within the boundaries of the present range. The Asiatic group is known from the upper Miocene, and throughout its entire history was much more widely distributed than in the present epoch,
latter.

forms of the

and

in particular, the

range included not only several parts of Asia,

present
range,

lying

outside

its

but

also

Europe.

In

the

zoogeographical sense, and partly also the systematic, the Asiatic

forms must be considered as relicts. The family is divided into three subfamilies

American (Cynarctinae) and two contemporary Procyoniae, the American raccoons and Ailurinae the Asiatic raccoons or pandas. The total number of genera is 18, of which 8 are contemporary. The number of described species of contemporary American genera is quite great, however, most of them undoubtedly constitue only races and the total number of species is, probably, only 11 or 12. The subfamily of American raccoons, Procyoninae, includes

one

extinct

the genera: Bassariscus CJentinkia"**) (cacomistles; 2, possibly 3

species),

Procyon (raccoons, 2 species), Nasua

1

(coatis,

species),

Nasuella (mountain coatis,

bears,
1

species), Potos (kinkajou, or

monkey

species), Bassaricyon (1, possibly 2, species).

The subfamily of Asiatic raccoons or pandas, Ailurinae, conof the genera Ailurus (panda, or cat bear, 1 species) and Ailuropoda (bamboo bear or giant panda, 1 species). Species of
sists

the family constitute in

order. Equally with

all

5%

of the total

number of species of
family
is

the

hyaena and bears,

this

one of the

least specious families of carnivores.

912

In practical terms, a series of species are innocuous,

some may

be harmful to hunting economy, and individual species are considered valuable fur-bearing species or

game animals

(raccoons).

In the fauna of our country, indigenous representatives of the

family are absent.

One

representative of the raccoon genus, Procyon

has been acclimatized as a fur-bearing species (V.H.).

Appears
to contradict statement in previous sentence
in

**Misspelled "Yentikia"

Russian original

Sci. Ed.

Sci. Ed.

1373

Subfamily of American Raccoons

Subfamilia Procyoninae
Gill,

1872

Genus of Raccoons

Genus Procyon

Storr, 1780

1780. Procyon Storr. Prodromus meth.

mamm.,

p. 37.

Ursus lotor

Linnaeus.
Size moderate.
In general

the fox or,

more

appearance and body structure, the animal resembles closely, raccoon dog. Head broad, with sharp
tail fluffy,

muzzle, ears large, protruding from the fur and tapering,

not prehensile,
its

length comprises about half, or slightly more, of

body

length. Digits long, deeply separated, grasping.

Skull moderately elongated, with broad braincase and well-

defined crests. Hard palate extends far backwards

of total length of palate.

to 1/2 or 1/4

Dental formula

I-C-P M = 40.
3

14 14

Molars and

last

upper
flat

premolar broad, more or less defined quadrate form, with

not developed.

masticatory surface and low, sharp protuberances. Carnassial teeth

Os penis present, to a greater or lesser degree curved. Omnivorous predators, associated with forest regions, good
tree-climbers. Northern forms in north of range
fall

into winter sleep.

characteristically
to the northern

American genus, distributed

in

North

America
and

border of the range of the family (see

characteristics of family above,

to the extreme, or

and description of species below),

to

almost extreme, southern limit of range of

the family in South

America

southwestern Brazil, Paraguay,

northwestern Argentina and southeastern Uruguay.

According

to its systematic position within the limits of the

is

family, the genus

quite closely related to several others and

is

not related to the

number of those which show

particularly sharp
it

deviation. Both morphologically and biologically

is

considered a

generalized, apparently, a less specialized group of the family.

1374

Within the limits of the genus, quite a

described. Seven of

lot

of species have been

them have been accepted at the present time by several authors. However, it is quite obvious, that there are in the genus 2 in all the North and Central American P. lotor Linnaeus, washer-raccoon, and the South and Central American P. cancrivorus Cuvier, crab-eating raccoon. These sharply distinguished species are sometimes relegated to different subgenera

{Eupro-cyon

for the southern species).

One

of the species (northern) belongs

among

the

most impor-

tant fur-bearing animals of

North America.

During the last decades, P. lotor was introduced into several European countries and into the USSR for acclimatization as a
quality fur-bearing species (V.H.).
913

WASHER-RACCOON, OR AMERICAN RACCOON

Procyon (Procyon) lotor Linnaeus, 1758
1758. Ursus
lotor.

Linnaeus.

Syst.

Nat.,

ed.

X,

I,

p.

48.

Pennsylvania^.

Diagnosis

Only species

in the

genus

in the

USSR.

Description
In

general

appearance

relatively

short

trunk,

short

legs,

generally stocky build and dimensions

and

in part in several char-

acteristics of color, the raccoon, most of all resembles the raccoon dog, and, to a lesser extent, the red fox. Tail relatively short and constitutes about half the body length. Head short and broad with a short pointed muzzle and quite large eyes. Ears set wide apart, large and protruding strongly from fur, with moderately

acute

tips.

Winter fur long and fluffy, quite soft, with a thick underfur. Tail covered with long erect hairs and appears thick and fluffy.

Upper surface of hands and feet clad in very short hairs their pelage differs greatly from that on legs. Callosities on lower surface of paws bare.
^Further

synonyms

very numerousrepresent nothing of

interest to us

and are

not provided.

1375

side,

General color tone of dorsal winter fur dull gray on the upper with greater or lesser admixture of brown, sometimes reddish

or chestnut hairs (mainly on nape).

Dark

tips of hairs

form

slight

wavy

pattern against general fur color.

On

lower surface, general

color tone slightly lighter; guard hairs here shorter and fewer and

underfur more obvious. General tone of underfur grayish-tawny. Fore legs and thigh region have same color as lower body surface, hind legs higher*; their heels are blackish. Hands and feet lighter,

almost white.
rings,

On

tail

are 5 to 7 sharply defined black or

brown

914

between rings (they are broader than rings) grayish or brownish. On lower surface of tail, dark rings not so sharply defined, their color lighter than on dorsally, and they are sometimes interrupted. Top of head gray. On facial portion, black or nearly black (brownish) "mask" sharply defined in form of large spot beginning on cheeks and extending, becoming wider, across eye to muzzle. Along upper surface of muzzle, it extends to lower part of nose forming narrow projection, which also extends backwards to forehead. Sometimes, stripe extending along muzzle slightly separated
and
its tip is

black. Color of parts

from eye spots by a slight brightening [see Fig. 346]. Facial mask is demarcated by well-defined white fields bordering it above and extending backwards under ears and to sides of neck. Lateral parts of muzzle, lips, chin and lower surface of lower jaw white. On
throat, a transverse black or

brown

area, separated dark

mask

which extends backward from muzzle. Ears clad with short grayish or brownish hairs and bear small black spots of varying size and definition at base of posterior surface. In addition to the raccoons of the described normal color,
pattern;

narrow white

stripe

black animals are met with (melanists).

There
nous,

is

one molt per year.

Facial region of skull short and quite wide. Braincase volumi-

obviously

swollen

from behind,

long

and

wide

in

interzygomatic region with nearly parallel borders. Facial length of

skull less than cranial. Nasal

the upper skull profile convex^

profile

line

gently

bones short and quite broad. Line of frontal region highest, from which descends both backward and forward.

Supraorbital processes very weakly defined, frontal surface between them somewhat concave longitudinally. Mastoid processes

Meaning

unclear; hind legs longer?

Sci. Ed.

1376

913

Fig.

346. Washer-raccoon, or American raccoon, Procyon lotor L. (Sketch by

A.N. Komarov).

Hard palate extends behind end of toothrow to approximately 1/3 of total palatine length. Auditory bullae rounded in form, inflated. Sagittal crest weakly
large;

paroccipitals moderately developed.

defined; occipital

well [defined]. Canines relatively short, upper almost straight, lower clearly curved. Fourth premolar and first molar of upper jaw almost equal to each other (molar a little larger),


1377

second molar approximately twice less than lower molars almost equal in dimensions.

first. First

and second

is 435-545 mm, of females, 410-570 200-405 mm, of females, 192-340 mm; length of hind foot (with the claw) of males is 96-138 mm, of females, 83-129 mm. Condylobasal length of male skull is 94.3-125.8 mm, of fe-

Body
tail

length of males

mm;

length of males

is

males, 89.4-115.9

mm,
fat

of females, 58.3-81.2

mm; zygomatic mm.

width of males

is

60.2-89.1

Weight

fluctuates very greatly, particularly in connection with

accumulation before winter hibernation

from 5.4

to 15.75

kg

and may even reach 22.2 kg^ (V.H.).

Systematic Position
In comparison to the second species of the genus
P. cancrivorus,

the described form

is

sharply characterized. In

some

respects,

it is,

apparently,

more specialized (V.H.). Geographic Distribution

Central and North America. Acclimatized in the

USSR,

Central

Europe and on several islands of the West

916

Indies.

Geographic Range outside the Soviet Union

Natural range occupies North America and Central America, south-

wards
is

to the

Panama Canal

zone.

The northern border of

the range

quite complicated

starting at the

lower reaches of the Saint

Lawrence River (near Quebec [city]), it extends along the south of Quebec and Ontario (Canada) provinces to Lake Nipigon (a little north of Lake Superior), starts along the shore of Superior, going
along
it

to its western extremity.

Thence,

it

directs itself to the

southwestern extremity of Lake Winnipeg, and farther, rises in a

curved line northwards to the western part of Lake Athabaska. Thence, the border sharply descends to the south-southwest to

information on dimensions and weight of raccoons acclimaand they do not give a correct idea about variation in these characters. Figures given above are from American sources and characterize variability of the species in its native land in general. Size and maximum weight after Hall and Kelson (1959), remaining data on weight after Burt and Grosenheider (1952).
'There
is

very

little

tized in our [country],

1378
the southwestern corner of Alberta Province. Farther, the border

takes a completely curious form.

It

describes a large extension, a

loop as

is

were, to the south, bypassing a significant (eastern) part

915

Fig. 347. Skull of washer-raccoon, or

American raccoon, Procyon lotor

L.

1379
of the Rocky Mountains and Great Basin and descends to 40 and even 35 N. lat*. From this region, being drawn towards the Pathe border passes northward, sometimes near it (southsometimes strongly distanced (northern), to the southeastern corner of British Columbia and, rising somewhat to the northwest,
cific

Ocean

ern),

reaches the coast of the Pacific ocean

at the level

of the northern

extremity of Vancouver Island.

In the Pacific Ocean, the range includes Vancouver Island and the small islands of Las-Tres-Marias along the Mexican coast. In the Atlantic Ocean, it is encountered on small islands lying directly off the coasts of South Carolina, Georgia and Florida.

916

Fig.

348. Natural range of Procyon lotor L. Somewhat simplified. Habitats in the Archipelago of the Windward Islands are not shown (after Hall and Kelson, 1959).

*The species range now includes most of

this

region

Sci. Ed.


1380
Generally speaking, raccoons are absent on islands of the West Indies; however, they are found on some of the Bahamas (in par-

Providence near Nassau'*), on Cosumel Island near Yucatan, and on the Lesser Antilles Islands, in particular, on
ticular,

New

917

Guadeloupe and Barbados (possibly also others) from the Windward Islands. The raccoon was acclimatized and lives in many parts of the F[ederal] R[epublic] of G[erman] and the G[erman] D[emocratic] R[epublic]. It was not introduced in other countries. Transgressions to Holland and northwestern France are known (Niethammer,
1963).

Geographic Range

in the Soviet

Union
raccoon
in

The experiment
began
in

in acclimatization of the

our country

1936, and was repeated more than once in various parts

number of attempts (up to 1962) was 26 1222 individuals. Of these, 64 were from the farms and zoos (38 imported from western Europe), the remaining were derived from a population which had been put together in the Trans-Caucasus (Zakatalo-Nukhinsk valley in Azerbaijan; Rukovskii, 1963). The animals were introduced in various places, often far from each other, and they never formed a significant, continuous range. Far from all of the transplantations (with respect
of the country. The total
releases, for a total
to the purely biological side of the matter)

had the same

results. In

several places, the animals died, in others, the results have not yet

been sufficiently determined or are unreliable, in several, the animals apparently survived well, obviously settled down and formed quite numerous populations^. In the Far East, raccoons (black) were introduced on Petrov Island near Vladivostok and a series of places in Primor'e territory (southern half), but all introductions were unsuccessful the animals died (on the map, these places are not shown; Abramov,

The suggestion that raccoons appeared (were introduced?) on the Bahama was based on a misunderstanding P. maynardi was described as early as 1898 from New Providence Island. 'Concerning information given below on places and results of transplantation of raccoon in our country are based on the most recent, in part unpublished, data and describe the situation in 1962 and the beginning of 1963. They differ significantly from Kolosov's (1958) and Rukovskii's (1960) communications, in which the success
Islands in 1932 (Niethammer, 1963)

of acclimatization was strongly overstated and already did not conform with the actual
situation.

1381

917

Fig.

349. Localities of introduction and occurrence of the washer-raccoon or American

raccoon, Procyon lotor L., in the

or results
unclear,
dots,

USSR.

Places where acclimatization

was unsuccessful

are

designated by crosses; places where the raccoon apparently

districts in

survived
stable

by

and the

Byelorussia and in Azerbaijan, where

are

more

or less

populations

were

constituted,

designated

by

blotches

(V.G.

Heptner).

1963). In Middle Asia, raccoons were introduced into former

918

and apple forests of Shan Mountain Range facing towards the Fergana valley in Arslanbob (north of Dzhelalabad). They were noted here in the Ach' region (between Arslanbob and Dzhelalabad) and in the Bazar-Kurgan region (west-northwest of Dzhelalabad; introduced in 1936 and 1952; Novikov, 1956). In January 1963 here "raccoons are practically absent"^. Raccoons were introduced into the mountain forests of the valley of the
Dzhelalabad
district of Kirghizia, in the nut

the southern slopes of the Tien

Pskem River (near kishlak* Sidzhak tween the Ugamsk and Pskem ranges
of Tien Shan; Salikhbaev
*P.

northeast of Tashkent, be-

the extreme western spurs

et al.,

1963).

Sagaraev, Board of Forestry and Nature Protection belonging to the Council

forest farms; their location

of Ministers of the Kirghiz Soviet Socialist Republic. Apparently, these introductions

(in

Chavachin and Karaamin

is

imprecise), and Yanushevich

(1963) also spoke of them, believing that these introductions were not successful,

recommending cessation of raccoon introduction * Local word for village Sci. Ed.

in

Kirghizia.

1382

According to one datum of Salikhbaev et al. (1963), the raccoon survived here, and has somewhat broadened the area of its
occurrence; according to other data

"the

results. ..were unclear",

is

and evaluation of
in

its

situation and

its

future fate

too optimistic

the literature^.
in

On

the whole, attempts to acclimatize the

their results as

raccoon

Middle Asia have been unsuccessful or

of 1963 are not clear and, apparently, they must be stopped.

Trans-Caucasus (Azerbaijan), introduction of the raccoon into the forest zone associated with the southern slope of the Main Caucasus range, beginning in 1937, was successful and led to the creation of a quite considerable and stable populaIn the eastern
tion (annual catch of

1000-1500

individuals), occupying one quite

from it. The main from the region of Belokana City (a little east of Lagodekha) along the so-called Zakatalo-Nukhinsk forest (Zakatalo-Nukhinsk valley) to Nukha city, and farther, to the Ismailla region, i.e. for an extent of more than 200 km. This region is the largest focus of occurrence of the species in the USSR, from
significant section, and isolated centers separated
strip

range extends as a

whence

is

taken material for future settlements.

is

separate section of occurrence

found

in the so-called

Kuba-

Khachmas forest massif from Kuba toward the northeast and north to Khachmas and Khudat on the railway line near the sea shore. At the beginning of the 60' s, this section was apparently not yet connected with Zakatalo-Nukhinsk.
in

small section of occurrence lies

the Talysh and

Lenkoran

forests.

On

the whole, in

1963

in

Azerbaijan, the raccoon inhabited Belokansk, Zakatal'sk, Kakhsk,

Nukhinsk, Vartashensk, Kurtkashensk, Ismaillinsk, Khachmassk and Lenkoransk regions (Ya.K. Aga-Zade, Nature Protection Board of
the Council of Ministers of the Azerbaijan SSR).
In the

ently, survived, in the riparian forest of the

North Caucasus, the raccoon was introduced, and apparTerek and along the
in the

Sulak [river]?

lowlands of Dagestan (south of the eastern

Cis-Caucasian lowland). Raccoons were introduced along the "left tributary of the Kuban'" (Aliev, 1963), but clear data about them
are absent.

The attempt

to settle raccoons in

Kabardin-Balkariya
to accli-

(black raccoons) was unsuccessful.

As shown by an almost ten-year experiment, attempts

matize the raccoon
in

Byelorussia were, apparently, quite success-

^M. Isamukhamedov, Board of Forestry and Nature Protection of the Council of

Ministers of the Uzbek. SSR.

1383

where, as a result of three introductions (52, 37 and 38 individuals in 1954 and 1958), a significant population was formed (on 1 January 1963, about 700 individuals). It occupies a section
ful,

along the Pripyat' River from Petrikov up almost to Pinsk (about

120 km), approximately 30

km

in

breadth along each bank

situation of the population

(Petrikovsk and Zhitkovichsk regions of Gomel' sk district, and

Stolinsk region of Brest district^).

The

has been evaluated as satisfactory.

Information in the foreign literature (Niethammer, 1963) concerning the occurrence of raccoons in Latvia (in spring 1961, 7900
is completely untrue. In Latvia, the raccoon-dog widely distributed, but the raccoon was not introduced at all. 919 The perspective on raccoon acclimatization in our country,

individuals!),

is

if

the Caucasus

mainly
is

the Trans-Caucasus

is

not considered,

Trans-Caucasus,
studied.

very limited from the purely biological side,

side has also not yet been analyzed (for

although steps already undertaken have not yet been sufficiently

The economic

details, see section

"Biology") (V.H.).

Geographic Variation
American authors, even
variation of the raccoon
the
is

most

recent, believe that geographic

very extensive.

Some

accept 30 races of

the species P. lotor, and another 4 species (insular) besides P.

cancrivorus, accepted here (Miller and Kellog, 1955); others recognize 24 races of P. lotor and another 5 species (one with two
races), besides P. cancrivorus (Hall

and Kelson, 1959). These coninto our country.

siderations are, apparently, exaggerated.

It is

not

known which forms were introduced

As

said, part

were derived from the zoological parks and fur farms

(V.H.).

Biology
Population. Experiments in acclimatization of the American raccoon first began in 1936, when the Uzbek Zoo introduced 22 raccoons in former Achin region of the Kirghiz SSR. In 1949, their

number here did not exceed 80-100

5-8 animals per 1000 hectares.
'A. Abaturin, State
isters

individuals, with a density

Committee

for the Nature Protection of the Council of

Min-

of Byelorussian SSR.

1384

An experiment in introduction of two pairs of black raccoons on Petrov Island in Primor'e territory in 1937 proved unsuccessful. In 1941, 21 raccoons were introduced in Ismaillinsk region,
Azerbaijan

SSR

(Rukovskii, 1951). In 1950,

it

was established

that

they already inhabited a series of regions in an area of 1300-1400

km^. The total number of raccoons reached 2000 individuals, with

density averaging
15.3 animals per 1000 hectares. In

1950, 44

raccoons were introduced into the Khuzakhsk and Khasav"yurtsk regions of Dagestan. In 1951, 28 raccoons were introduced 70-90

km

to the south of

Krasnodar

city.

In

1952, 33 raccoons were

introduced into the former Oktyabp'sk region of Kirghizia, and 79

raccoons into the Zakatal'sk region of Azerbaijan SSR. In 1953, in former Khizinsk region of Azerbaijan, 17 raccoons were introduced, while 43 raccoons were settled in former Bastandaksk in Kabardinsk region of Kazakhstan, and 16 black raccoons

A[utonomous] S[oviet]

S[ocialist] R[epublic] south of Nal'chik in

50 raccoons were introduced in Petrikovsk region of Gomel' sk district of B[yelorussia] SSR, 55 raccoons in former Budennovsk region of Primor'e territory, and 104 raccoons on Zelenchuk river in Stavropol' territory. In all, from 1936-1954, 534 raccoons were
the region of the Nal'chik and Belaya rivers. In 1954,

introduced for acclimatization.

From

1941,

all

introduced animals,

except 16 black raccoons, were caught valley of Azerbaijan (N.N Rukovskii).

in the

Zakatalo-Nukhin

In 1954, 486 raccoons (Abramov, 1961) caught in the TransCaucasus were introduced in Partizansk, Anuchinsk and Yakovlevsk regions of Primor'e territory. The introductions were accomplished
in

coniferous

broad-leaved

forests in the valleys of

montane

taiga

and forest resources were insuffew, and food were with hollows fires, trees there, tree lowlands, but ficient. The raccoons went off to the hollows were also few, and the raccoons settled in burrows, clefts
rivers. In these forests there

was extensive

tree-felling

of rocks and haystacks. They were not able to accumulate fat before winter and they spent the winter badly
(its

duration

is

120-150

days) with severe frosts. Mortality was recorded among them from hunger and cold. In following years, the population everywhere

was

less than in the year of introduction.

locally, but mortality

was

greater. In the 60' s, the

Reproduction occurred raccoon became

extinct in Primor'e.

1385

920

Ryabov, 1963), raccoons settled in a narrow belt with an area of about 7000 km^, extending from Novorossiisk to Lazarevskaya. Here, they were sporadically encountered everywhere in low density. In better places in Goryache-Klyuchinsk region 3.5 animals per 1000 hectares; in 0.1. In more 0.6; in Lazarevsk region the Gelendzhiksk region favorable lands of the Ismaillinsk region, density reached more
In Krasnodarsk territory (Kotov and

than 20.0 animals per 1000 hectares (Rukovskii, 1951). In their native land, in boggy lands of Wisconsin state (USA), density of

raccoon settlement fluctuated from 25.7 to 34.6 per 1000 hectares (Dorney, 1954). Under more favorable conditions of the southern
states,

raccoon populations

may

attain a very high density.

Up

to

1949, their highest density was considered to be established in 111.0 animals per 1000 hecIllinois; in an area of 384 hectares

tares, but in the state of Missouri, 102 raccoons were caught in an area of 25 hectares; i.e. here their density was 408.0 (Twichell and

Dill, 1949). In the state of

Ohio

(Butterfield, 1944), in

open hunt-

ing lands, were established densities of: 36.1; 142.0; 166.6; for
reserves

340.0

(66 raccoons on 193.5 hectares).

Habitat. In Kirghizia, raccoons inhabit montane forests domi-

nated by walnut trees. In the Trans-Caucasus, they prefer valleys

with broad-leaved forests consisting of oak, white-leafed poplar,

smooth-leaved elm, alder, walnut and chestnut. In these forests, there are many lianas, wild almond-leafed willows, hops and hol-

low

ests almost

Raccoons no less willingly settle in montane beech fordevoid of undergrowth (N.N. Rukovskii). one of the few wild In the USA (Nelson, 1930), raccoon animals, whose populations increases with cultivated appropriation of territory. With free water and forest vegetation, raccoons
trees.

here settle also in the

swamp

lands (Dorney, 1954); the presence

of trees of large diameter with hollows are preferable but not

necessary.

rous predators. In composition

Food. The American raccoon belongs to the group of omnivoits food includes a variety of both

animals as well as plants. Their combination, and the ratio of each, depend upon the biotope, time of year, and the yield of each food
(see Table 71).

In the northern

USA,

in

swamp

lands of the state of Wisconsin

(Dorney, 1954), the raccoon's main food consists of crustaceans, fish, young muskrats, insects, grapes, fruits and berries (see Table
72).


1386
Table 71. Seasonal characteristics of raccoon nutrition
occurrence. Becker, 1945)
in

Texas (%

of

Type of food
Birds

Spring

Summer

Autumn

Winter

Snakes
Fish

Freshwater crustaceans Mollusks

Wasps
Beetles

4 4
18

23
21

22

Crickets

Larvae and pupae of butterflies

Spiders

Water scavengers, and diving

beetles

Water bugs Acorns Wild grape Mulberry

Persimmon Hawthorn
921

55 14
occurrence)

16

16

10
7

47 38
2

29 24

5")

10
3

36
6

1
13
('

Table 72. Seasonal characteristics of raccoon nutrition in Wisconsin

Food type
Adult muskrats Young muskrats
Cricetid rodents

III-IV

V-VI
6

Vl-VII
2

IX-X
2

X-I
12

20

Adult ducks

Duck eggs
Eggs of other birds
Fresh water
crustaceans
Fish

10
49
20
7

40
11
1

35
2'

9
1 1

3
1

12

4 52
17
1

64
22
5
11

64
3

Frogs
Insects

39

16

25

27

Maize Acorns
Grapes Wild pear Raspberry

50

60

10
19

14

The frequency of young muskrats in the raccoon diet in various swamplands fluctuates from 8-9 to 19% (Dorney, 1954). In the state of Washington (Tyson, 1950), the occurrence of mollusks in the diet of adult raccoons in summer was 85%, of crustaceans 85%, of fish 65% and of neris worms* 50%. In the stomachs of

Meaning

not clear; probably refers to marine polychaetes

Sci. Ed.

1387

fish

young raccoons, mollusks constituted 67%, crustaceans 100%, 45%, worms 11%, milk 78% of the contents (20 stom-

achs of adults and 9 of young).

In the Trans-Caucasus (612 excrement samples; Rukovskii,

1951, 1957); the raccoon feeds mainly on the following kinds of

food (% of occurrence):

Mammals
Birds

5.5

Fish

2.3

11.7

Crabs
Insects

2.6

Amphibia
Reptilia

52.6
13.2

82.4
12.9

Mollusks
Plants

63.9

Therefore, under

new

conditions, the raccoon

shows

itself to

be a typical polyphagous
habits.

animal, like the

badger as regards food

Of mammals, it feeds on mice and voles; of birds mainly on small passerines and injured ducks. Of reptiles, raccoons eat Trans-Caucasian frog. turtles, lizards, rarely snakes; of amphibians Among insects were encountered beetles, orthopterans,

lepidopterans, dragon flies and caddis flies,

among them

a significricket,

cant

number

fall

among

pests of agricultural plants

mole
fruits

locusts, crickets, long-horned beetles, bark beetles, click beetles,

leaf beetles, weevils

in the

and others. Raccoons feed also on plant foods:

of lianas

Trans-Caucasus, most frequently wild grapes,

and acorns. To a lesser extent, they eat pears, apples, cherry plum and blackthorn. In spring and winter, raccoons
(sasparilla), nuts
utilize

more animal food,

in

Home

range. Dimensions of the

summer and autumn home range of

plants.

the adult racits

coon are associated with the abundance of food within

borders

and the distance of feeding areas from shelters. In Azerbaijan

(Rukovskii, 1951), the area of such a home range fluctuates from 4 to 9 km^ with a radius of 1.0-1.5 km from shelter. A lactating female does not travel more than 600 m from her litter. Consequently, the area of her home range does not exceed 144 hectares. In the state of Ohio (USA), during two years, no cases were recorded where a raccoon traveled more than 1.5 km from the

922

was marked (Butterfield, 1944; according to recapture of 91 marked raccoons). On average, the animals were caught no farther than 400 meters. Burrows and shelters. In Azerbaijan, the raccoon settles in hollows of white-leaved poplar trees, rarely oak (Rukovskii, 1951,
place
it


1388
1955). This

is

explained by the faci that hollows of size suitable

for raccoon, are here

more frequent

in poplars.
is

Average height of
is situ-

hollow trees inhabited by raccoons

ated at a height of 10-12

about 21.0 m, their circum-

ference (at breast height) reaches 4.5 m. The living cavity

m from the ground.

cm

The average diameter

Amplitude of daily temperature fluctuation inside such a cavity does not exceed 3.5C, and in hot summer hours, the temperature is 10-12C lower than that of the outside air. Besides the main living hollow, within the home range of the raccoon, there are also temporary shelters hollows, where the raccoon spends the day resting. In North America, raccoon also settles in caves and among rocks and clefts. In the absence of tree holes, it settles in earthen burrows, muskrat huts, boxes for nesting ducks and other shelters (Dorney, 1954). During the capture of 297 raccoons in the state of Ohio, only 8% of them were hidden in tree holes, 47.5% among rocks and in burrows, the remainder under roots of trees, on newly fallen trees, etc. Of 316 raccoons, 46.4% occupied ground burrows, 18.2% tree hollows, 14.4% shelters in the base of trees, and 10.0% among rocks. Raccoons occupied 22% of the artificial
of the entrance hole
is

19

(not less than 12 cm).

bird nests of the

box type (Butterfield, 1944). Raccoons often occupy burrows of marmots, sometimes in open places. Litters were also found in haystacks and heaps of brushwood. It is supposed that the raccoon was initially a species closely

associated with tree hollows.

As

a result of cutting

down

large-

trunked trees containing hollows, and the transformation of forest

areas into agricultural ones, the raccoon

became an animal capable

is

of living under various conditions of cultivated landscapes.

Daily activity and behavior. The raccoon

turnal animal.
It

a crepuscular-noc-

emerges from
at

its
It

hole at sunset and sometimes

it

returns to
its

it

even

day break.

climbs trees well, but

gathers

food mainly on land

along

river banks, in
it

swamps and

other

similar places. In searching for food,

sometimes rambles through

is

shallow water and along sand bars.

characteristic feature of the raccoon

it.

that

it

rinses

its

gathered food in water before eating

"'poloskun", (= to

Hence, the raccoon's name

wash off)Hibernation and winter sleep.

In districts with

snowy, cold

winters, the raccoon, as well as the

brown bear and badger, have

a characteristic winter sleep. In the Trans-Caucasus (Rukovskii,

1389
1951), in association with

warm

winters and brief frost, raccoon

It emerges when the soil is and Canada, where snow lasts for

activity stops only during snowfall.

partly free of snow. In Kirghizia

a longer time, raccoons leave their shelters earlier, before the

snow

thaws

in spring. In

Canada, their sleep

is

lasts for

about four months

of the year. Here they are able to survive hollows while the temperature of the outside air

down

to

-43C.

Seasonal migrations and transgressions. After the introduction of 250 raccoons from Ohio into Arkansas, their appearance

was noted

37, 75 and 255

km

from the place of introduction

(Butterfield, 1944).

Reproduction. In Canada, i.e. in the northern parts of the range America, estrus occurs is March. In yearling females, estrus takes place in May-June. In the state of New York, raccoons mate at the end of January-beginning of February (Mackeever, 1958).
in

In the state of Michigan, the majority of females

beginning of March, and few of them

of June.
In the states of Florida

at the

mate in Februaryend of March; indi-

vidual cases of mating were observed in Virginia at the beginning

tion lasts

from 10 February
in

females are mated, in

in

and Georgia, the season of reproducAugust: in February, 14% of the March 41%, in May 12%, in June 12%,
until 3

July

12% and

August

6%

(Llewellyn, 1953*). After a preg-

923

nancy lasting for 63 days, raccoons bring their young into the light in the period from April to the beginning of October. Practically half of all [parturition] occurs in May. The number of embryos averages 3.2, fluctuating from 2 to 5. In the state of Michigan, the average number equals 4, with fluctuation from 3 to 7.
In the Trans-Caucasus, young appear in April-beginning of May. The quantity of newborns fluctuates from 3 to 8, more

frequently 4 (Rukovskii, 1951).

Growth, development and molt.

length of the

embryo reaches

and on the 46th day 65 mm (Llewellyn, 1953)**. Young are born blind, and eyes open on the 20th day. At 1.5 months of age, they cut all teeth. At the age of 2.0-2.5 months, they already accompany their mother while searching for food, and at 4-5 months old, they switch over independent foraging. In September-October,
*Not in Lit. Cit. Sci. Ed. **Not in Lit. Cit. Sci. Ed.

On the 21st day of pregnancy, 24 mm, on the 35th day 45 mm,

1390

young raccoons do not

Raccoons
Sex

yield, in terms of dimensions, to adults,

but are markedly lagging behind them in weight.

attain sexual maturity at the

age of about one year.

ratios in populations are different: in the states of

Illinois, the ratio

favored females

animals, even 14:86), in the state

i.e.

46:54 and 40:60 (among hunted of Missouri (62:38 and 61:39,

Ohio and

the ratio favored males).

Out of 102 raccoons caught in an area of 25 hectares, 32 were alone, and in 12 cases caught in pairs; in 6 cases the raccoons were obtained in trios. Only one time were groups of 4, 5 and 6 raccoons were caught. The animals were caught in winter during winter sleep (Twichell and Dill, 1949).
Enemies, diseases, parasites, mortality, competitors and population dynamics. Wolf, lynx and eagle-owl are regarded as

enemies
these

of the raccoon. In places inhabited by raccoons in the

predators are rare and do not cause great losses to
it.

USSR,

There are no data on diseases and mortality of raccoons. In Trans-Caucasus, competitors of the raccoon in obtaining food are shown to be brown bear, badger, otter, pine and stone martens, jackal, cats, wild pig,
red deer and fat dormouse (Rukovskii, 1951, 1953).

There are no data on population dynamics. In Kirghizia, the raccoon population increased 5-fold within 13 years, and in
Azerbaijan

100-fold
it

within 9 years.

Field characteristics. The crepuscular-nocturnal activity of the

raccoon makes
Their presence

very difficult to carry on direct observations.

the characteristics of inhabited

may be known by

hollow (claws marks on bark, polishing of entrance into tree-hole, etc.) and by the tracks of their paws on damp soil along the banks of water bodies. In the track of the anterior paws, prints of five
long clawed digits are clearly seen. Tracks of the hind paws resemble, in strongly

reduced aspect, tracks of the hind paws of brown

bear.

They

are

more extended

in

length than in the badger

(P.Yu.).

Practical Significance

The American raccoon

mainly used
in the

is

a valuable fur-bearing animal. Its hide

is

manufacturing of men's winter fur coats. In

view of the

fact that the districts of its probable distribution should,

apparently, be restricted to the zones of

mixed and broad-leaved

1391

forests, the practical significance of

raccoon as a fur-bearing ani-

mal cannot be

great. Its practical

importance

siderable in the southern regions of the

fur manufacture are few.
lands, they

may be more conUSSR, where objects for

may cause

losses to

With occupation by raccoons of marshy young muskrats. Raccoons do not

pose any danger to adult muskrats and aquatic birds since they only feed on injured individuals. Raccoons are useful as they destroy insects harmful to forests and agriculture. (P.Yu.).
924

Family of Mustelids Family Mustelidae Swainson, 1835

AMERICAN MINK'
Mustela (Mustela) vison Schreber, 1777

nil?
1949. 1958.

Mustela vison Schreber*. Die Saugethiere .... pi. 27. Eastern Canada. Mustela vison tatarica Popov V.A. Tr. Kazansk. Filiala

AN

SSSR. Seriya

biol.

s-kh. nauk, 2, p. 135. Tatari.

i

Lutreola vison altaica. Ternovski. Biologiya

akklimatiz-

atsiya amerikanskoi norki (Lutreola vison Brisson) na

Altae (Biology and acclimatization of the American mink

...

in Altai), p. 15.

Middle course of Sary-Koksha River,

Altai. (V.H.).

Diagnosis
Color and skull are same as in Russian mink, Mustela (Mustela) lutreola, but upper lip has same color as top of head (not white) and postorbital constriction of skull is sharper its width is less

than width of interorbital area. Dimensions

'In

somewhat

larger (V.H.).

sometimes called the "eastern mink". There is new, completely artificial name. This name also the range of the American mink, with respect to the European, lies is confusing nearer to the west, in the western hemisphere. ^Some of our monographers (Novikov, 1939, 1956; V. Popov, 1949; Temovskii, 1958) refer to Brisson as the author of this name with reference to 1756, and they
our literature,
this species is

no basis for the introduction of

this

try to designate the priority of this assignment to Schreber. This is a clear misunderstanding. Zoological nomenclature begins in 1758 (publication of the 10th

even

Edition of "Systema Naturae" by Linnaeus) and

all

names given earUer

are generally

not

accepted.

*Mispelled Schreder

in

Russian original

Sci. Ed.

1392
Description^
In general appearance and construction, the American mink is completely like the Russian; however, the tail is on average somewhat longer, and reaches or almost reaches half the body

length (from 38 to 46 and even 51%).

Winter fur
longer,
fur
is

is

as in the Russian mink, but denser, close-fitting,

more luxuriant and softer. As fur, its value is higher than of the European mink. Length of contour hairs (Altai) on back

27.7

mm, on
is

belly

is

22.2

mm,

guard hairs are 22.7 and 16.6

mm,
1

underfur

14.8 and 10.6, respectively.

is

Number

of hairs on

cm^ on sacrum
According

19,530; on shoulder, 21,783 and on the belly

17,320 (Tserevitinov, 1958).

to

other

data,

in

Altai

(Ternovskii, 1958),

number of

hairs

on

mink (4 specimens) cm^ in middle of back

ranges between 22,664 and 37,100, consisting on average about

22,458. For one guard hair, there are from 31 to 34 underfur hairs. Length of guard hairs on back is 20.9 mm, of underfur hairs, 12.3

mm.
There
is

one pair of inguinal

an exception,

teats,
six.

and three pairs of abdomi-

nal, in all eight; as

General tone of winter fur tawny of various intensity, from

very dark blackish-tawny to light-tawny. Color
uted over
all
is

equally distrib-

of body, lower side only slightly lighter than upper.

Guard
part very
is

hairs, bright,

dark-tawny, on spine often approaching

black, straight and elastic, but not coarse. Underfur hairs on dorsal

925

wavy, grayish-tawny with a bluish tint. Base of underfur and closer to tip, intensity of dark tone strengthens, and sometimes end of tail has pure black color. Sometimes, along the back, a broad wash of somewhat darker stripe is displayed. Chin and lower lip white white color does not spread onto upper lip as a rule. Color exhibits quite significant individual variability, which
light bluish-gray. Tail darker than trunk,

may be somewhat

different in populations comprising various parts

of our country. This applies to both qualitative and quantitative

aspects (for cause, see section "Geographic Variation"). Percent-

age ratio of different saturated color types varies

'All data given below, especially

in

different
to

where no individuals are preserved, belong

populations from natural conditions within the

scarce and fragmentary.

USSR. Best known

are populations in

the Tatar Republic (V. Popov, 1949); Altai (Ternovskii, 1958); in others data are very

1393
populations, but dark and very dark animals predominate. Thus, in
Tatariya, tawny-black animals constitute

48%, dark-tawny 36%, tawny 10% and the light-tawny 6%. Frequently on skins (in Tatariya, about 24%) white guard hairs ("hoary") are encountered and sometail tip is

times the
In

white

(in Tatariya,
is

about 4%).

some

individuals, underfur

white

in

color and

may
and

occupy from 40 to 80% of skin uted on the entire body surface.

surface of the body
lie

area, or they

may even be
lower

distriblip

In addition to white

chin, very often there are irregular white marks noted on the lower

and groin. They form of patches and sometimes stripes. The number of patches and the area they occupy vary. In the Altai, 29.2% of them were on the lower lip and throat, in 30.4% they extended from lower lip to fore limbs, in 35.1% they extended from lower lip to anus, in 4% they were
throat, neck, chest, belly

on

along the mid-line of the animal venter

in the

distributed on the groin, in

lower

lip,

throat and groin,

0.1% between fore paws, and in 0.1% on chest and

in

1.2% on
It is

groin.

noteworthy that the white marks may sometimes (3.3%) not occur

925

Fig. 350.

American mink, Mustela (Mustela) vison Schreb. Sketch by A.N. Komarov.

1394
at all (data

from 727

skins).

As

a rule, the upper lip in the

in the Altai, in

Ameri-

can mink

is

never white. However,

36.4% (213
Their di-

skins of 585) white patches occurred on the upper

lip.

mension varies from weakly defined

half the
lip.

to those

occupying more than

in other re-

In Tatariya, this

is

absent. In the Altai, the white

marks

are, apparently,

more strongly developed than

gions of our species range.

the neck and chest,
is

broad white patch (as

in

marten) on

rarely encountered in the Russian

it

mink M.

(M.) lutreola; in our American mink,

has not been mentioned.

is

Considerable development of white patches

characteristic of

cage minks.

It is

very interesting that in Tatariya, with naturaliza-

tion of the animals, within 10-11 years, the

number of

skins with

white patches on chest, belly and groin, decreased sharply. At the

end of
926

this period

(1938-1948), only animals with small white

lip,

patches and only on lower

chin and throat were found (V.

this

Popov, 1949). Apparently, there was a gradual elimination of

feature.

which the ends of the guard hairs are twisted, similar to singed hairs, are rarely met with. This "singing" is a mutational deviation known from caged animals in the Altai (7.8%). On the whole, color variation of American minks inhabiting the
in

The animals

USSR

is

much

greater than in Russian [mink].

in color

Sexual differences

and
is

in

development of patches and

hoariness are absent. The latter

males (Tatariya) or
approximately 2

in

more often developed, either in females (Altai, 94.6%). White underfur

is

apparently occurs more often in females. Hair length in females

mm

less than in

males (middle of back, V.A.

Popov, 1949).

mink has not been described in from winter, but is shorter, sparser and dull. Length of summer guard hairs on dorsum is 17.3 mm, and 9.8 mm; on 1 cm^ of dorsum there are about that of underfur 17,450 hairs on average, for 16-22 underfur hairs for each one
fur of our [American]
detail. Its color differs little

Summer

guard hair (Altai).

The
massive.

skull is similar to that of the

It is

European mink, but

is

it is

more

narrower and less elongated, cranium

wider, shorter

and more flattened. Postorbital constriction is more sharply defined in narrowest part, its width is greater than width of skull between orbits. Zygomatic arches, especially in middle portion, are thinner and weaker than in Russian mink, but diverge somewhat

1395

more widely. Mastoid processes well-developed and fuse with lower part of occipital crest. Auditory bullae are flat and broad; in posterior part they are set apart from each other not so far as in European species. Ratio of their length to their breadth approximately 1.0:1.5. Protuberances,
crests, etc. of skull are generally

more sharply-defined than in Russian mink; occipital crest developed significantly more strongly; sagittal crest well[-developed]. Upper molar larger and more massive than in Russian mink, its

area, in frontal projection,

no smaller or only

slightly smaller than

its

area of upper carnassial tooth; longitudinal diameter of

well

developed inner lobe more than logitudinal diameter of outer lobe. Posterior end of second upper premolar more or less strongly transgresses into notch found at anterior end of carnassial tooth. Inner

surface of main cusp of lower carnassial tooth with well-developed

sharp

rib.

On

the whole, in

its

general appearance, specialization

of skull as that of a predator skull more strongly developed than

in

Russian mink, and as regards degree of specialization

skull.

itself,

somewhat reminiscent of polecat

Skulls of females, besides general dimensions, distinguished

by somewhat less development of crests, protuberances, etc. and are somewhat lighter. Skulls of young are characterized by the more rounded and expanded braincase and weakly developed
facial portion.

Geographic variation of the skull within the borders of our

country not expressed (see beyond).

The OS penis of

the

American mink, having a

structure typical

of the genus, differs considerably from that characteristic of the

European mink, not only in size, but also in form. If one looks at the bone from below, its terminal part is not deflected to the right and the bone is curved in one plane. Its base is laterally compressed and is significantly wider along the vertical. Basal third is also flattened. Along middle of sides, there is a short groove, and on lower surface of the anterior part, there is a deep groove (Novikov, 1956; V. Popov, 1949). There are 14 thoracic vertebrae, 6 lumbar, 3 sacral and 18-21 caudal (40 individuals). Weight of heart in males (16) is 5.3 M8. 4-1 1.5 g, in females (9), 4.5 M5.46.18 g; cardiac index
is

7.25-M9.71-12.79. Length of intestine

in

males

mm

1635-M 1806-2 140 mm, in females, 1220 M1480-1690 (Altai). Number of caudal vertebrae in Tatariyan mink is
is

19-22.

1396

927

Fig. 351. Skull of

American mink, Mustela (Mustela) vison Schreb.

1397
928

Diploid number of chromosomes

is

30.

is 340-450 mm, of females, 310-375 156-247 mm, of females, 148-215 mm; length of hind foot in males is 51-70 mm, in females, 40-58 mm; length of ear in males is 20-29 mm, in females, 19-26 mm (in 110 males and females from Tatariya and 41 males and 31 females from Altai). Weight of males in winter (Altai and Tatariya) is 500-1580 g, of females, 400-780 g. Condylobasal length of male skull (32; Altai") is 62.1-M66.472.6 mm, of females (14), 57.0-M59.3-61.6 mm; zygomatic width of males is 35.0-M37.1-41.4 mm, of females, 31.2-M32. 8-34.4 mm; mastoid width of males is 30.3-M32.8-36.8 mm, of females, 27.6-M28.5-30.5 mm. Os penis length (24) is 35.9-M40.5-45.4 mm. Weight is 0.110MO. 229-0. 580 g. Age differences in dimensions and particularly weight of OS penis are very significant. Length of bone in young (30; Tatariya) is 37.9-M41.7-45.2 mm, weight is 0.130-M0.1670.210 g; in adults (30; Tatariya), length is 39.9-M44.6-49.7 mm, weight is 0.240-M0.336-0.450 g (V.A. Popov, 1943). Difference

Body
tail

length of males
is

mm;

length of males

in

weight

is

double.

in body measurements and weight is very Measurements of females (Tatariya) constitute 82-85% on average of measurements of males, while weight is 60-80%. Weight is very strongly variable according to season, reaching its maximum in autumn. At that time, males in Tatariya weigh about 1000 g, maximum 1580 g, females about 600 g, maximum 780 g. Least weight of males happens from April to June, females from June to August (nursing young). Average weight of the animals, and in part measurements, obviously change by year (V.H.).

Sexual dimorphism

significant.

Systematic Position

According

to

morphological characteristics, the American mink

is

closest to the European.

These concern

ethological characteristics.

main ecological and Some authors (among the most recent

its

examples, K. Zimmermann, 1959; Gaffrey, 1961) even consider it as only a subspecies of the European, and suggest (Gaffrey,
1961), that in Middle Europe, remaining natural populations of the
"For skull dimensions of Tatariyan mink, see section "Geographic Variation"

1398

European mink disappears (extirpated) due to its hybridization with the American, dissolving it so to speak, into the population of the
latter.

According to our data, however, uniting the two minks into one species, cannot be accepted. They are close ("paired"), but independent species. This is indicated not only by their morphowhite logical, and most of all, craniological, characteristics (color marks on lips, a feature which is not very stable), but also by the

absence of hybridization in nature here, in part a result of asynchrony of the reproductive cycles. It is possible that there is
also a lack of compatibility in genital systems also occurs (see
structure and size of os penis). Sharp antagonism exists in nature between these species not only competition, in which the larger and stronger M. (M.) vison crowds out the European mink, usually simply destroying it (V. Popov, 1949; V. Popov et al., 1954). Accurate material data on hybridization of the two mink species in nature are absent, apparently not only in our country, where

this

Their experimental crossing

question was especially studied, but also in Central Europe. in captivity was also unsuccessful
yields hybrids with the black polecat [M. putorius] (see above

(Schmidt, 1933; V. Popov, 1949). At the same time, the Russian

mink

and below). The American mink itself represents a sort of further development of the European mink a further stage in the specialization towards predatoriness. This is indicated by the structure of its skull,

with constricted interzygomatic area of braincase, more strongly developed protuberances and crests of skull, stronger dentition,
etc. In

comparison with the

skull of the

American mink,

the skull

of the European bears several infantile features.

929

The "paired" mink species show

a quite remarkable parallel-

ism with the "paired" species of polecats black M. (P.) putorius and white M. (P.) eversmanni. The European mink corresponds to to the second. This parallelism is the first, and the American

manifested

in a series

of features

first

of

all,

in skull structure

and

its

interzygomatic region, relatively wide

in the less special-

ized black polecat and constricted in the white. Features of paral-

lelism in skulls of polecats with both pairs of "mink-polecat" are,

however, developed at a higher level of specialization mink do not attain such a height as each corresponding polecat species.
This
is

the parallelism of a pair of terrestrial species, in part of


1399

even desert-steppe species [polecats], and a pair of species associated with the aquatic environment. In this situation, the level of
specialization found in the polecat cannot be attained, possibly
related to smaller and

weaker prey

[of mink].

The

significant quantity of shared characters with polecats, on

American mink) and on the other with kolonok does not allow us to split off mink as a separate subgenus, much less a separate genus, and also separate polecats {"'Putorius'") at a generic level from remaining species of genus Mustela. It is
the one hand (especially
also hardly acceptable to separate mink from the genus Mustela and to unite them, as a subgenus, with an independent genus of

polecats
link

Putorius (Stroganov, 1962).

Mink

serve as a connecting

between polecats and other species of the genus Mustela, especially the pair European mink black polecat. They also produce hybrids. The American mink itself represents a branch of this trunk, with white polecat its summit. (V.H.).

Geographic Distribution
Beginning with 1933, the American mink, usually
both in
its

in fairly large

groups, were introduced into very different parts of our country,

the Caucasus.

were introduced, but quickly for further establishment, wild animals from populations naturalized to various places, began to be used, in
start,

European At the

part, in Siberia, in the

Far East, and also

fur farms

in

captive

mink from

particular, in the Altai. After the first years, introduction of ani-

mals within the range of Russian mink was stopped, or almost new groups of mink were directed mainly into Siberia and the Far East where the Russian mink is absent. At the beginning of the 60' s, and still earlier in some places (Altai), fairly
stopped, and

considerable areas occupied by American mink were formed.

However, in the majority of cases they were isolated from each other and one continuous range of this species is still absent in our country. There is a basis for considering that, on account of the new introductions and the natural dispersal, it will spread over
time, although
species.

some

territories are

poorly suited as habitat for this

As a result of absence of sufficient data on distribution of the mink in separate locales, we will tabulate below only districts where it was introduced, irrespective of how many times and in which


1400
parts

of these

regions

the

animals

were introduced. Also,

is

intraregional distribution of local populations

not considered.

Starting in

1933, American minks were introduced into the

931

European part of the USSR in Voronezh district (first experiment Voronezh preserve). From 1933 to the beginning of 1963, the mink was introduced in various quantities to Murmansk and Arkhangelsk districts, Karelia, in Kalininsk, Gork'ovsk, Volgogradsk, Chelyabinsk and Voronezhsk districts, and into Tatarsk, Bashkirsk, Mariisk as well as Lithuanian and Byelorussian Republics. Beyond the Urals, animals were introduced in Sverdlovsk, Tyumensk, Omsk, Kemerovsk, Novosibirsk, Chitinsk and Irkutsk districts, in Altai and Krasnoyarsk territories, in Tuvinsk, Buryatsk, and Yakutsk Autonomous Republics, into Magadansk, Kamchatsk and Amursk districts, into Khabarovsk and Primorsk territories, into Chukotsk national regions and in several other places, among them Sakhalin
and Urup Island in the Kurils. In the Caucasus, introductions were in northern Osetiya, in Georgia and Azerbaijan, and in Middle Asia in the Tien Shan on

territory of Kirgiziya.

The most northerly place of introduction Kola Peninsula, lower Severnaya Dvina, mouth of Ob' River, mouth of Nizhnaya Tunguska, Podkamennaya Tunguska (middle course), Vilyui, upper

Kolyma and lower Anadyr'.

In the expanse

between the Pacific

Ocean and upper and middle courses of the Ob', mink are established generally southward, extending or almost extending to the
state border. In western Siberia
try, it

and

in the

European part of coun-

extends southwards to the steppe zone. On the whole the range is not continuous, and separate places of introduction and separate regions of acclimatization, one may say are still isolated

from each

other, in

introduction in

great distances. Regions of Asia are particularly Middle and the Caucasus

some places by

remote from other places.^

932

were introduced These introductions have not yet given results in the sense of creating sufficiently significant populations from the
In a series of places, small groups of animals
in a

few

spots.

perspective of being considered permanent. This applies to the

^Materials used in the construction of the

map and

the considerations given were

obtained from the Main Board of

Farm, of the Council of Ministers of RSFSR (Chief N.B. Eliseev; L.S. Tamantseva). Some data were obtained from V.A. Popov and G.A. Voronov).

Game

1402

931

Fig. 353. Natural species distribution area of the

American mink, Mustela (Mustela)

vison Schreb. (from Hall and Kelson, 1959).

'

Extreme North, northwestern and in part western parts of the European USSR, as well as the Caucasus and Middle Asia. More positive results were apparently obtained in Byelorussia. Quite complete and numerous populations were created in the eastern European territories of the country in Tatariya, in part in Bashkiriya and other adjacent places. Here populations are of exploitable number and density. Everywhere, in regions of

habitation of the indigenous mink,

i.e.

to the

west of the Ural, the

American mink has crowded out and exterminated it. In Siberia and the Far East, where in several districts living conditions are favorable for the introduced species, two quite sig-

1403
nificant populations, both in

numbers and

in territory occupied,

already have spread out.

One occupies

the Ussuri territory, includplaces. In Trans-

ing Priamur'e, and the other

Altai and adjacent

Urals (western Siberian lowland) where living conditions are less

favorable for the American mink, numerous attempted introductions did not succeed.
It

is

evident that a significant number of

introductions into Siberia, in particular in the Extreme North and

northwest, did not lead to the formation of stable and entirely

naturalized populations.

Geographic Range outside the Soviet Union

(natural).

The mink occupies

the greater part of North America.

The

north-

ern border extends along the northern shore of the continent, excluding, however, the

from Hudson Bay

the tundra

to

Ungava Peninsula. Moreover, in the expanse Amundsen Gulf in the Arctic Ocean, the
at

northern border extends approximately along the southern limit of

from Eskimo Point

Hudson Bay

to its intersection

of

the northern coast of the continent at 125 west long.

to

The western border is formed by the Pacific Coast southwards San Francisco and the eastern by the Atlantic Coast, except

the eastern extremity of the

Nova

Scotia peninsula.

The southern

border extends along the coast of the Gulf of Mexico, approxi-

mately to Corpus Christi, and thence

line across the states of Texas,

is

directed along an irregular

New Mexico

Utah, Nevada and

northern California passes to the Pacific Ocean.

Mink

are absent

on the Aleutian and other islands of the Bering Sea, on Kodiak, on the Queen Charlotte islands and several other small islands along
the Pacific Coast of America. In the east, the mink is absent in Newfoundland and the islands of the Gulf of Saint Lawrence. In Europe the acclimatized mink is found in France, in Central Europe (F[ederal] R[epublic] G[ermany] and the G[erman]

D[emocratic] R[epublic]),

is

widely distributed in the Scandinavian

it

Peninsula. In Cent[ral] and in part in west[ern] Europe,

has

replaced and destroyed the European mink (V.H.).

Geographic Variation
Very many geographic forms of mink have been described from
the natural range in North America. Recent authors accept 14 races


1404

and moreover, a separate species M. macrodon extinct (exterminated) about 1860 and inhabiting,
vison,
apparently, an extremely limited territory in the northeastern United
States

of the species M.

(New Brunswick and

state of

Maine). The actual number of

is

races of the species in America, apparently,

less,

and M.

macrodon

differing

only in somewhat larger dimensions hardly

deserves to be

split off as a separate species.

were settled by captive-born mink from various large fur farms. The greater
In the beginning, the various parts of our country

part of the material consisted of animals born in captivity in our

farms. Animals directly imported from outside our borders constituted only a minority.

The

origin,

i.e.

the races to

belonged, even

if

we assume

that at least a part of

which this material them were wild

933

unknown. However, they were all, apparently, captives. mink in the USSR proceeded by means of animals captured by us from the wild. During study of the results of acclimatization, several real characteristics of the Tatarsk and Altaisk mink were established, and separate subspecies were described tatarica and altaica (see synonymy). Difference between both "forms" lies in dimensions {tatarica is larger) and degree of development of white marks (in altaica, they are more, and may even appear on upper lip) and density of pelage (in altaica, it is slightly denser). The relationship of these "forms" to American mink is unclear. The separation of the mentioned "subspecies" has no basis, in that what is signified by the word "form" does not correspond to the concept of subspecies. The word applies to very young populations, only yet being established from genetically heterogenous material {tatarica mainly from Pushkino near Moscow, Kola fur farm, 1937). At the time of their study, 1934; altaica they had existed in all for 14-15 years. Both populations themselves bore features of their "domestic" origin, in the form of
mink,
is

Later, introduction of

occasional mutations that are

unknown

in wild

mink, or are ex-

tremely rare (albinism, variant of underfur color, "scorched").

From published
is

material (V. Popov, 1949; Ternovskii, 1958)

it

were subjected to the action of intensive selection and were rapidly changed general dimensions and skull measurement changed, as well as frequency of occurrence and size
clear that populations

of white marks, etc. All of this

is

entirely natural for animal groups

it

with a heredity "uncoupled" during cage raising;

cannot be used

1405
as a basis to accept the

lished geographic races in

above-mentioned populations as estabthe present sense of this word. One can

only speak about differences in these artificially created populations.

In the meantime,
selves,
it

is

entirely unclear

how

they will form them-

even

after several decades, especially in case of

union of

ranges, not to mention the time needed for elaboration of actual

subspecies.

Dimensions of mink from Tatariya (110 males and females) at from the Altai (41 males and 31 females), at beginning of the 50's (Ternovskii, 1958) are the following: body length of males from Tatariya is 375the end of the 1940' s (V. Popov, 1949), and

M417-450 mm,
Altai of males
375

of females,
is

mm;

tail

length

330-M358-371 mm; the same from 340-M395-450 mm, of females, 310-M345of males from Tatariya is 195-M224-247 mm,

of females,

178-M 195-2 15 mm; tail length of males from Altai is 156-M184-205 mm, of females, 148-M158-173 mm; length of
is

hind foot of males from Tatariya

males, 50. 0-M54. 5-58.0
Tatariya
the

6 1.0-M65. 0-70.0

mm,
Altai

of feis

mm;

the

same of males from

51-

M60-67, of females, 40-M50-55 mm; length of

is

ear of males from

22-M26-29 mm,

of females, 2 1.4-M22. 8-26.0

is

mm;

same of males from

Altai

20-M22-25 mm,
mink
skulls

of females,

19-

M21-23 mm.
Average
measurements
0.31

of

from

Tatariya:

condylobasal length of males (20)

(19), 60.65

is 68.95 0.47 mm, of females mm; zygomatic width of males is 39.75 0.40 mm, of females, 33.9 0.23 mm; interorbital width of males is 15.38 0.31 mm, of females, 13.2 0.17 mm (for craniological

data on Altai mink, see above in section "Description") (V.H.).

Biology
Population.

From 1933

to 1948,

about 4000 mink were introduced

into the territory of the

USSR,

distributed on

50

sites in

22

oblasts.

To

1964, the total population of animals introduced into the Euro-

East,

pean part of the USSR exceeded 2000, and in Siberia and the Far 14000 (V. Popov, 1964). Concerning the results of these introductions one may judge that in 1957-1959, in the RSFSR (excluding Yakutiya) from 26.8
thousand wild minks of both species were caught. Of this number, American minks did not exceed 20-25%; i.e., the number

to 35.7

1406

.^v

4y,.

934 Fig. 354. Arkhara river a place of introduction and occurrence of the American mink in Arkharinsk region of Amursk district. June 1956. Photograph by A.G. Pankrat'ev.

of American minks caught per year was from 5350-6450 to 7150-

8925. Separate count of the yield of each species were not made;
therefore, the ratio of their

number

in regions

where both species

existed can only be estimated very approximately. Moreover, at the

present time, the figures on skin preparation reflect not just the
934 population

numbers of the species but the numbers of hunters,

spend
in this trade, etc.

their skill, the time they

Due

to the strength

of a series of causes, mink, in

many

districts, are

hunted less than

would be allowed by

their population.

More

than

80%

of the American minks are obtained from

Siberia and the Far East, mainly from the Altai, Krasnoyarsk and

Khabarovsk
try

territories.

From

a series of indirect data,

it

can be

estimated that the total number of the American mink in our counapparently does not exceed 30-40 thousand.

1407
Habitat. With respect to choice of habitat, there are no differ-

ences between the American and European mink. In Georgia,

the Alazan' valley (Ekvitimishvili, 1951),

in

around forest creeks 1.0-1.5 m steep banks freezing only along the
1955), lands favorable for

mink stations* are areas wide and 15-100 cm deep, with

sides. In the Altai (Ternovskii,

and have a good network of spaces under the shallow ice. Banks are well protected, steep and covered with trees containing a large amount of fallen limbs and branches, good undergrowth, with an abundance of mouse-like rodents. There, 3-5 mink are usually found along 1 km
are rich in fish,

mink

of shore-line. Satisfactory lands can be distinguished from less

favorable conditions for nesting and protection; banks are weakly
forested and with poor cover.

On one km

of shore-line, only 1-2

mink

are encountered in such an area. Unfavorable areas are poor

in fish,

or

if rich,

then there are no under-ice cavities and few

open water

areas. In

one

km

of shore-line in these conditions, no

first

with. Surroundings of the and second types predominate in the lower courses of small in the upper reaches of rivers. ers, and the third

more than one mink may be met

riv-

Food. Of greatest significance in foods of mink in Tatariya (V. Popov, 1941, 1949) are voles (36% occurrence), fish (28.8%), crustaceans (26.7%), frogs (17%) and aquatic insects (19.8%).
935

Importance of each food changes noticeably according to season. In winter, foods obtained from water prevail: fish (45.5%), crustaceans (32.2%) and frogs (17%); the percentage of voles decreases
to

10%.

In spring, the significance of terrestrial animals increases;

occurrence of voles increases to 55.6%, of insects

comprise

6%

to 20.7%, birds and occurrence of crustaceans decreases to 18%; in

food remains, small hares are encountered. In summer, a greater diversity of food is characteristic. Vole occurrence is high (40.6%),
crustaceans (30%), insects (29.6%), frogs (21.2%), birds (mainly 16.7%. In autumn, voles in the form of nestling) and their eggs

occur

in

50%

of data, fish in

15.5-38.7%, and crustaceans

in

22.4%.

Within the boundaries of montane Altai (Ternovskii, 1955), in American mink, mammals constituted 58.5% (occurrence); rodents 53.8% (including voles 50.2%), and shrews and
the diet of the

moles 4.7%. Birds were met with in 3.3%. Reptiles constituted 1.0%, amphibians 5.8%, fish 54.9%. Among birds, 11 different species were recorded most frequently, dippers {Cinclus cinclus]

*The Russian word

is statsiya;

stantsiya might have been

meant

Sci. Ed.

1408

and pine grosbeak [Pinicola enucleator]. Among fish, small ones prevail: minnow, gudgeon [Gobio gobio] and wide-headed sculpin [Cottocomephorus], in all, 11 species of the 26 species known in Altai. Mollusks constituted only 0.05%, insects 25.6% (mainly from stomachs of fishes). Here also (Berger, 1947), in the earlier period
of acclimatization, the occurrence of insectivores in the food of

fish

10.2%. 41.8% of mink food, water voles25%, 12.5%, fish 20.9% and insects 12.5%. (Vladimirov, murid rodents 20.5%, water voles 30.1%, 16.9%, 15.7%, amphibians 1.2%, and 14.4%.
insects
constituted
birds
In Irkutsk district

American mink constituted 3.9%, hares mouse-like rodents 43.6%, water voles

5.1%, 14.1%,

squirrels
fish

2.6%,

7.7% and

In Sverdlovsk district (L.M. Tsetsevinskii), mouse-like rodents

1940),

constituted

birds

insects

In connection with differences in the composition of aquatic

and bank fauna, a well-defined geographic variation is observed in the foods of American mink. The absence or scarcity of river crustaceans in

many

of the water bodies of our country

is

particularly

evident. In the Far East, in the food of

are

American mink, crustaceans replaced by fresh-water amphipods which are eaten in great

quantities (Yu.A, Salmin and V.D. Shamykin). In the first years

American mink, living for a series of generaon farms, food obtained on land predominated, but feeding characteristics quickly changed character, and approached that of
after introduction of

tions

wild mink

in type.

America (V. Popov, 1949), warm-blooded animals have greater significance in the food of mink, than in the USSR
In North

(mouse-like rodents, muskrat).

The mink

eats

4-9 times

daily.

The quantity of food eaten

of the live weight of the ani-

during this time constitutes

20-25%
The

mal. In Tatariya (V. Popov, 1949) and in the Altai, the American

mink makes

a store of food.
1

size of the store reaches

1

1200 g
1

(6 frogs, 3 water voles,

tundra vole,

viper, 9

minnows and

wide-headed sculpin) (Ternovskii, 1955). Home range. Dimensions of the home range of each individual is subject to seasonal change. In mink having a settled mode of
life,

area of the

home range

in

southern Tatariya on average equaled

about 16 hectares, fluctuating from 0.1 to 40.0 hectares (V. Popov,

1949). Dimensions of daily range of activity in winter fluctuate

from

0.1 to 2.1 hectares. In

summer, when food yield increases,

1409

^*

936

Fig. 355.

Brood

shelter of
district.

American mink. Arkhara

river in

Arkharinsk region, Amursk

June 1956. Photograph by A.G. Pankrat'ev.

the
is

home range

decreases; usually only a small part of the range

exploited, sometimes, only a radius of

in

50-80

from the bur-

row. In the Altai,

winter mink live in an area of 200-300 m, not

wandering more than 100 m from the bank, and are confined to open water (polynia) (Ternovskii, 1955). In the Far East (N.M. Berger), 0.8-1.0 mink may patrol on 1 km of the shore-line, in 0.04-0.10; in 0.3-1.0; in Krasnoyarsk territory eastern Siberia

Kuznetsk Alatau 0.7-0.8. banks ments along the 7-10 km. autumn up
river
to

In spring, the animals undertake

to a distance of

up

to

move20-30 km; in

936

In

America (USA)
is

in

an area of 440 hectares (280 hectares of

which

dry land), 17 minks were captured and tagged (14 males

1410

and 3 females); 5 of them (29.4%) were caught a second time in the same winter. During the following 4 years (winters of 1945/48) in the same area, only 6 new minks were caught, out of which 2 were repeatedly caught (4 times). Males were considerably more active than females and the area of 440 hectares was insufficient for them; one male, in the course of three years moved on to the area every month, making a round trip of about 1.5 km in one and rethe same route and then returned. Females were less active peated hunting was achieved within a distance of from 25 to 400 m from the place of marking (McCabe, 1949). Burrows and shelters. In the Volga-Kama territory, there are 4 types of burrows of American mink (V. Popov, 1949). They are constructed frequently in the hollows in root masses of growing

and rarely in hollows of fallen ones. The holes are often masses of hummocks ("koblakh")* of the alder which are typical for flood-land alder swamps. These holes are characterized by a large number of entrances and twisting passages. Rarer are earth holes of structure similar to that of the European mink. The number of exits of the mink burrow varies from 1 to
trees,

in the root

8.

In the breeding burrow, exits are often stuffed with grass so

young cannot crawl away. Usually, the breeding burrows are situated not more than 20 m from the bank of the water body. Burrows of single animals are also encountered (considerably more rarely) at a distance of up to 50-60 m. The bedding of the nesting chamber consists of dry grass and leaves, wool and underfur of consumed animals and wood dust. Out of 24 mink shelters investigated in the Altai (Ternovskii,
1955),
937

54% were

found

in

hollows of fallen
trees,
soil layers.

trees,

34%

hollows among the roots of openings among rocks and

with water,
at

4% in hummocks
Mink

in intraroot

and

4% in

shelters, in the latter

case, are found in sections of floodlands

2-10

from the water

it. The bedding in Carex sedge, pediformis ("puzzle grass"). Daily activity and behavior. The mink is active at various times of day. The rhythm of diel activity changes in different seasons in relation to security of food, prevailing means of obtaining it, weather

from

which are not submerged no more than 50 the nesting chamber usually consists of
level, but

conditions, etc. Thus, in the rut period, animals are highly active

*Local word?

Sci. Ed.

1411

throughout entire 24 hr period. For the period of pregnancy, daily

activity typically

shows two peaks.

In the period of nursing young,

the female

is

active most of the 24 hr period. In

summer, the

litters

are active mainly at twilight. Later in autumn, the activity during the day light hours again increases (Ternovskii, 1955).
In Tatariya, in

American mink

in winter (January) three peri-

ods of activity are observed with an interval between the peaks of

approximately 8 hours (V. Popov, 1949). The

period, lasts

first

a daylight

from 10

until 14 hr

and

is

associated with obtaining

food

in

water bodies. The second

nightfall.

at twilight,

begins

at

18 hr,

somewhat before
decreases.

The

third occurs in the

second half of

the night and the pre-dawn period. During severe frosts, activity

The mink is maximally active in July-September, and November-February. Males are more active in March and less from the end of rut until August. Females are maximally active in July-September. In winter, in a 24-hr period, the mink moves from some tens of meters to 4-5 kilometers. In the Altai, the mink is most active morning and evening, and
minimally
in

also at the end of the night (Ternovskii,

1955). In winter and

spring.

summer,

activity

sion occur in spring, and in autumn, dispersion of

searching for

Rut and disperyoung and adults, home ranges, and then mink are maximally active. In
is

less than in

autumn and

cloudy calm weather, activity grows. In winter, with

ture

air

tempera-

from -20

to

-37C, the mink

is

active for not

more than 51-

64 minutes during the 24 hr period. In December-January, it leads a hidden, subnival mode of life. If in October and the first half of November, the length of the 24 hr trail reaches 800-1000 m, then during the period of deep snow, the daily trail decreases by 3-6 times and does not exceed 100 m. During swimming, undulating movements of the trunk play the main role. The swimming web is no more developed than in ermine, weasel and kolonok. In warm water (24''C), mink can swim

more than 3 hours without stopping. In cold water, animals died within 27 min. During this time, body temperature fell from 40C to 19-20C. With water temperature up to 8C, the body of the mink cooled down under water for 118 minutes, while in ermine and forest polecat in 26-28 minutes (Ternovskii, 1955).* Seasonal migrations and transgressions. In winter, the character of activity in mink depends upon the abundance of food and its
for

*The mink body

is

better insulated in water

Sci. Ed.

1412

938

Fig. 356.

Tracks of American mint; on fresh new-fallen snow. Fudzin

river,

western

slope of Sikhote-Alin'.

December 1956 (Photograph by A.G. Pankrat'ev).

availability.

Complete freezing of water bodies, absence or small

air

number of polynias,*
under ice
call forth

holes and other opportunities to penetrate

emigrations and sometimes considerable conin those places

centrations of

mink

where food

is

more abundant

and available. In areas of the meadow type with little food, some minks lead a migratory mode of life, performing movements of 2-3 or up to 4-5 km in a 24-hr period in an area of floodland of 60 hectares (in the presence of abundant food, minks do not move away from the burrow more than 50-100 m). Migratory animals restrict themselves to one place for several days, and then perform a movement to settle down in a new place, again for several more days. Young mink, dispersing, also undertake migrations. More significant translocations were observed in the first period after introduction of mink, especially in cases of up
to
*Ice-free water surface

Sci. Ed.

1413
unsuccessful choices of places.
tion site a distance of 15

Mink moved from

40-60

the introducin Siberia,

km
to

(Tatariya) to

km

and for a distance of up

938

100-120

km

in the

Far East.

Reproduction. In our country, estrus in American mink proceeds from the end of February to the beginning of April. For the

most
1941,

part,

mating takes place

i.e.

in

mid-March

(Kler, 1941); (V. Popov,

in

1949),

month

earlier than

Russian mink. Estrus

separate individuals lasts one month, with 6-8 day intervals.

The

percentage of fertilized females equals 95-100. Duration of preg-

nancy
in

is

51.3 days on average, fluctuating from 40 to 65 days

(Kler, 1941). Differences depends

embryonic development, which

lasts

on duration of the latent period from 2-3 weeks to 1 month.

After formation of the placenta, development goes on for 25-26

Embryonic mortality is great and reaches 60-70%. It is most determined by magnitude of litter and depends upon external conditions of environment which control condition of the female organism. Birth takes place during the period from 4 to 22 May. In the Altai, the first signs of spermatogenesis were noticed in January. Already by 10-15 March, all stages of spermatogenesis, and sperm were in the tubules of the epididymis. On 1 February, pre-estrus proliferation of the uterus was observed in the female (Ternovskii, 1955). The first signs of rut were noticed at the end of February. It proceeded throughout March and the first half of April; peak of rut the second ten days of March. Its general duration here is 45 days. The number of young in the litter fluctuates from 1 to 9, and in exceptional cases, reaches 11 (Tatariya) and 16 (USA). In southern Tatariya, average litter size fluctuated in various years from
days.

of

all

3.2 to 5.8 (V. Popov, 1941).

Growth, development and molt. Weight of new-borns fluctu-

from 6 to 11 g. They are born blind, but develop rapidly, and eyes open at age of 30-31 days. Differences in increase in weight and dimensions between males and females are clearly noticeable only after 40 days. The lactating period lasts for 2-2.5 months. Mink's milk contains 3.8% lipids, 6.2% protein, 4.6% sugar and 10.66% mineral salts. Already at the age of 20-25 days, young minks begin to try food brought by the mother. In autumn, they attain adult dimenates

sions.

1414

On

farms, minks live 7-10 years, but are usually killed for

their skins before reaching their

age

limit.

Enemies, diseases, parasites, mortality, competitors, and population dynamics. The larger, stronger American mink ousts and
depresses the European.
939

There are little data concerning population dynamics, and these data are not always reliable. The history of establishment of mink in the Altai gives some conception of growth rate of the transplanted populations. Acclimatization was started in 1937. In 19371941, 248 minks were introduced. During the period 1940-1953, 431 minks were transferred from one region to another, to accelerate the transplantation process.

As

a result, during a period of 19

years, the

American mink

settled in

hundreds of rivers and streams

and occupied a

territory of

79,400

kml By

the beginning of the

1960's, they numbered several thousands (Ternovskii, 1955).

Field characteristics.

It is

not always easy to distinguish be-

tween the tracks of American mink and European. Jumping with paired paw prints is a typical gait for the European mink. Length of the jump is 25-35 cm. In American mink, tracks on the snow most often form a group of 3-4 paw prints. Paired tracks and the unique pace forming a chain of tracks are rarely observed. Length of a jump is significantly greater 55-60 cm. In the Altai, maximum jump length of the American mink is 1200 cm, while running, the mink can jump to a height of 50 cm.

It

can attain a speed of 20 km per hour for short distances; at a moderate walk 8-12 km, and in slow movement 3-5 km. The mink cannot run in loose snow deeper than 15 cm. Tracks of the paws of males are always larger than females; the male track is equal to 4 x 3.5 cm on average, but the female

track

is

3.5

x 3.0 cm. While moving slowly,

is

the length of the

female jump

always shorter than the male. Sex can also be

determined by the characteristics of urination on the snow. The weight load on 1 cm^ of foot surface is large, 15-31 g (average of

20 samples freezing on
otters, are

20
it.

g).

Weak

fur covering of the foot prevents

snow
as in

"Sliding hills" and grooves in

mink

trails,

explained by their need to dry the fur in winter after

coming out of water (Tarnovskii, 1955).

Identification of tracks and burrows of

mink may

also be de-

termined by contents of excrement and food remains near entrances

to

burrows (P.Yu.).

1415
Practical Significance

Acclimatization of the American mink in the

colonization in water bodies where
it

USSR

achieved

its

was previously absent for some reason or where it v/as destroyed. The American mink, as an object of the fur trade, gives a larger and higher quality skin than the Russian mink and its settling in water bodies, mainly in montane regions of southern Siberia and Far East, may increase the commercial yield to us of valuable "colored" furs. Trade in American mink in the USSR began in Tatariya in 1938. At the present time, the animal is mainly used for further settling by means of its capture in live traps. In some parts of North America, the mink destroys muskrats in
quite significant quantities, especially the young. There are analo-

gous complaints
both abroad and

in Siberia.

In past years, intensive breeding of the

in the

American mink

in farms,

USSR,

has pushed hunting of wild

mink

to

a secondary position (P.Yu.).

940982

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Works
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cited here are listed in both Parts la

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Part

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Mammals

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this

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Sci. Ed.

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990

Index of Latin

Names

of Taxa*^

aberrans

cf.

Meles
Mellivora

abyssinica

cf.

admirata

cf.

Mustela

adustus cf Canis

Aeluroidae 54
aestiva cf. Mustela

clamitans 383 Aonyx 492, 498, 854, 883 capensis 853, 854 853,854 hessica 883
cinerea 851,

anakuma cf. Meles Anurocuon 383 Anurocyon 383

Ailuridae 54

Arcticonus 404

Ailurinae 911

Arctocyonidae 54

Ailuropoda 54, 402, 907, 908, 909, 911

Ailuropodidae 54
Ailurus 54, 402, 907, 908, 909, 911
alhperakyi
cf.

erminea 663 663

tobolica

Arctogale 636, 640

Vormela
196,

Arctoidea 54

196 208 groenlandicus 208 208 208 lagopus 195, 196, 199, 202, 205, 206, 207, 208 208 semenovi 207 spitzbergenensis 206, 208 ungava 208
beringianus,
fuliginosus
halensis
innuitus

Alopex 62, 63, 68, 99, 194, 194, 195, 268 beringensis 196, 207

Arctonyx 493, 498, 817

arctos cf Ursus

arenarius cf Meles

Artiodactyla 10, 11, 54

asiatica cf.

Martes

aureus

cf.

Canis, Mustela

aurobrunnea cf Lutra
australis cf.

Dusicyon, Mustela

56, 63, 65,

196, 197,

pribilofensis

196,

baicalensis cf Ursus

196,

barang

cf.

Lutra

Bassariscus 69, 907, 908, 911

alpinus

cf.

Mustela, Kolonokus

Bassaricyon 911
beringiana cf Ursus
beringianus
cf.

altaica cf.

Mustela

cinerea 853

Amblonyx 498, 853

americana cf Martes

Ursus

birulai cf. Mustela,

Kolonokus

boccamela
bosniaca

cf.

Mustela

Amphicyoninae 62 Amphicynodontinae 62, 402 amurensis cf. Meles

Borophaginae 62
cf.

Martes Martes
the Russian origi-

brachyura

cf.

Reproduced from
nal appear in the

the Russian original. Page

numbers of

left-hand margin in the text

General

Editor.

'Page numbers with description are given in bold and those with illustrations, in
italics.

88

1540
Broiliana 817

brokmani

cf.

Mellivora

Caenolestoidea 9

canescens

cf.

Meles

Conepatus 498
Canidae 54, 55, 57, 58, 60, 61, 62, 69, 272,

402,907,911 Caninae 62, 63, 66

Canis 53, 62, 63, 68, 96, 96, 98, 99, 132,
194, 268

hodophylax 145 hungaricus 108 108 145 kamtschaticus kurjak 145 lagopus 196 141 lanka 108 100 383 108 lupus
indicus

itaUcus

24,

136

194,

laniger 123,

latrans 63,

lepturus
lupaster

56, 59. 63, 64, 96, 97, 100, 123,

99 1

adustus 99 albus 123,136 142 alopex 108 383, 386 arabs 145 argunensis 142 arundinaceus 139 aureus 108 63 = 53 141 146 beringiana bemardi 146 campestris caragan caucasica 00 caucasicus 108 chanco 142 communis 137 coreanus 24 corsac 265 crucigara 319 cruesemanni 108 cubanensis 140 145 desertorum 141, 142 dingo 61,95,99 142 dukhunensis 383 dybowskii 136 ecsedensis 108 100 gregoryi 146 140 143
63,
altaicus 139, 139,

123, 124, 125, 129, 132, 133, 135, 136,

137, 137, 138, 139, 139, 140, 141, 141,

31

algirensis

alpinus 63,

24,

123,

63, 64, 96, 100,

100, 101, 102,

105, 107, 108,

australis australis

antarcticus

bactrianus

bailey i

31

137,

123, 139, 140, 142

31

123, 141,
123,

24,

deitanus

123,

lycaon 146 mackenzii 145 manningi 146 marocanus 108 melanotus mesomelas 108 minor 108 montana 319 moreotica 108 nana 108 niger97, 146 24 145 pambasileus 145 primaevus 383 procyonoides 69 124,143 146 schensi 53 simensis 63 soudanicus 108 145 108 108 145 tundrarum 146 turuchanensis 136 vulpes265, 318, 319
137,

142, 142, 143, 145, 146

31

63, 99,

100,

99,
1

orientalis

pallipes 123, 141,

rex

rufus

signatus

syriacus

typicus 100,

tschiliensis 142,

24,

52, 53,

Canoidea 54, 55, 58, 402, 497, 498, 907,

911
caparioccoj
:
:
;

dorogostaiskii

24,

cf.

Mustela

124,

capensis cf Mellivora

Capreolus capreolus 837

Carnivora 10,
1

familiaris 53, 56, 96,

1,

49, 52, 53, 60,

907

caucasica cf Mustela
caucasicus cf Meles, Ursus
cavifrons
cf.

hajastanicus

24,

hattai 124,

Ursus

1541

Cetacea 10,

11

dinniki

cf.

Cephalogale 402

desertorum

Ursus
pallipes cf. Canis

Cervus elaphus 837

Dinocyon 402
Diprotodontia 9
dolichocrania
cf.

chamus
chanco

cf.

Fennecus
Canis
Lutra

cf.

Vulpes

Charronia 498, 502, 505, 508, 604

chinensis
cf.

dugong

Dugong

17, 23,

24

Chiroptera 10, 11

Chrysaeus 383

brachyurus
cinerea
cf.

Chrysocyon 62
59, 62, 63

Amblonyx
cf.

cinereo-argentatus

Dusicyon

63 culpaeus 63 hagenbecki 63
australis

20, 27, 24 Dugongidae 17, 23, 24 Dusicyon 62, 68, 268

cinereo-argentatus 63, 67

Clethrionomys rufocanus 521

rutilus

521

collaris cf.

Ursus
cf.

colore griseo

Vulpes

Conepatus 493, 498

consica
cf.

ecsedensis
Edentata 10
Eira

cf.

Canis

Mellivora

498

coreana

cf.

Mustela
Martes
Vulpes

Embrithopoda 23

coreensis

cf.

Creodonta 54
crucigera
cf.

culpaeus

cf.

Dusicyon

854, 863, 884, 885, 889, 891, adustus 392 384, 385, 386, 386, 388, 390, 391, 392, 392 dukhuneusis 392 fumosus 392 hesperius 392 infuscus 392 Enhydriodon 854 jason386, 392 Enhydris 882 javanicus 386, 392 992 Eomellivora 803 392 Eotheroides 24 392 Euarctos 403, 404, 406 primaevus 392 Eumetopias 53 sumatrensis 392 Eumustela 637
lutris

Cuon

60, 62, 63, 383, 383, 384,

386

alpinus 60, 63,

kamtschatica 884, 895 883, 894, 894,895 marina 894 895 895 883 895
nereis
orientalis

Enhydra 50, 492, 493, 498, 499, 854, 882, 882, 883 gracilis 895

reevi

stelleri

laniger

lepturus

Cuoninae 62
cylipena
cf.

Euprocyon 912
Eutheria 10

Mustela
euxina
cf.

Cynalopex 265
Cynarctinae 911
Cynodictis 60

Vormela
cf.

eversmanni

Mustela

Cynodon 402
Cynodontidae 402

Eyra 493

Cynaonyx 854

D
Dermoptera 10
Desmostyliformes 23
desertorum
cf.

Felidae 54, 55, 57

Canis

56 manul 50 56
libyca
sylvestris

Felis catus

56

1542
Feloidea 54, 55, 497

H
hagenbecki
Halicore 25
Haiitheriinae 23
cf.

chamus 63 63 63 zerda 59 zerdus 63

pallidas
riippeli

Fennecus 62, 68, 268

Dusicyon

Halyanassa 42

hamgyensis

cf.

Martes

Ferae 10, 49
ferghanae
cf.

Mustela

malayanus 403, 483

Helictis 493, 498,

Helarctos 397, 398, 400, 403

Ferungulata 10, 54
Fissipedia 53, 54,

817

60

heptneri

cf.

Meles, Mustela

flavescens

cf.

Vulpes

Foetorius 636
foetorius
cf.

javanicus 52
hoole
cf.

Heestes

auropunctatus 52

745

Vulpes

foina

Martes
Ursus

huli cf.

Vulpes
cf.

fontanieri cf. Mustela

hungarica

Mustela

formosanus
fuliginosus

cf. cf.

hungaricus

Alopex

Canis Hyaenidae 54, 55, 57, 60 Hyaenodontidae 54

cf.

G
Gale 636
Galictis 493,

Hyracoidea

10, 22,

54
25

Hydrictis 853

Hydrodamalidae

17, 23, 24,

498
Ursus

gedrosianus
Glires 10
glogeri
cf.

cf.

gigas
I

Hydrodamalis
25,

18, 23. 25,

25

27

stelleri

25

Mustela
Canis
Vulpes

Hydromustela 637

Grammogale 498
gregoryi
cf.

griffithi cf.

Grisonella 498

groenlandicus

cf.

Alopex, Ursus

Gulo 493, 498, 500, 501, 614, 614, 616,

albus 629 618, 628 618, 628 biedermanni 618, 628 borealis618, 628 gulo 617, 618, 618, 619, 622, 624, 628, 628, 629, 629 kamtschaticus 618, 629 katschemakensis 629 Iuscus618, 629 629 616, 618 615, 618, 628 vancouverensis 629 618, 628 wachei 618,628
arcticus

6] 7, 641, 803

boccamela 644 kamtschatica 644 644 644

nikolskij
nivalis

Ictis

636

Ictonyx 498, 783

inaurita cf. Mellivora

arctos

indica

cf.

Mellivora
cf.

indochinensis

Martes

58,

mesomelas

cf.

Canis

Insectivora 10, 11,

54
cf.

isabellinus-syriacus
itatsi cf.

Lutra

Mustela
cf.

intermedia

Martes, Vormela

luteus

schlosseri
sibiricus

japonicus
jessoensis

cf.

Ursus Ursus

vulgaris

cf.

Guloninae 641
gwatkinsi
cf.

Martes

gyas

cf.

Ursus

kalmykorum

cf.

Vulpes
cf.

katschemakensis

Gulo

1543
Kolonokus 673, 640, 690, 691, 723
alpinus
australis
biruiai

664, 686 700 664, 686 coreanus 700 686 700

raddei
sibiricus

kolymensis
koreana
cf.

cf.

Ursus

Martes
cf.

koshevnikovi

Vormela

kuatunensis cf Martes

kutab

cf.

Lutra

993

L
labradorensis
cf.

Ursus

Lagomorpha

10, 11

Lamprogale 502, 604

laniger cf. Ursus,

Lupus

larvata cf. Mustela

lasiotus cf.

Ursus

Latax 882
Leptarctinae 497

Leptonyx 785
leptorhynchus cf Meles

Leucocyon 194
leuconota
cf.

Mellivora

leuconyx cf Ursus
leucurus
libyca
cf.

Meles

cf. Poecilictis

884 868 incarum 853 kamtschatica 855, 867 kutab 869 854, 856, 858, 859, 862, 865, 866, 867, 868, 869 853 marina 884 867 minor 718 mira 853 monticola 868 865, 869 nudipes 854 oxiana 867 865 853 provocax 853 867 867 884 sumatrana 853 867 636, 640, 727 924 719 europaea 718 719 vison 924
gracilis

isabellinus-syriacus

855,

lutra 853,

855,

866,

maculicollis

meridionalis

855,

nair

855,

perspicillata 853,

platensis

seistanica 855,
stejnegeri 855,

stelleri

vulgaris 854, 855,

Lutreola

altaica

borealis

lutreola

lineiventer cf. Mustela

Lutrinae 497, 498, 817, 851, 853

Lupulella 99

Lutrogale 498, 853

124 124 karanorensis 24 123 124

altaicus
filchneri
1

Lupus 96

Lutronectes whitelyi 855, 863

laniger

tschiliensis

luscus cf
luteus cf.

Gulo Gulo

62 thous 62 Lycaon 63 thous 63

microtis
microtis

Lycalopex 62, 68

58, 60, 62, 63,

386

pictus 59, 60, 63

Lutra 493, 498, 500, 500, 501, 851, 851,

Lycaoninae 62
lymani
cf.

amurensis 867 868 annectens 853 aurobrunnea 855, 869 855, 867 barang 866, 869 canadensis 853 868 enudris 853 853
855, angustifrons
baicalensis

852, 853, 854, 854, 882, 883

Mustela

Lyncodon 493, 498

M
mackenzii
cf.

Canis

Mammalia 10

chinensis

Manati balaenurus 25
gigas 25
cf.

manchurica

Mustela Ursus

felina

mandchuricus

cf.

1544
marinus
cf.

Ursus Ursus

maritimus

cf.

Marsupialia 10

Martes 493, 498, 501, 501, 502, 502, 503,

504, 505, 507, 516, 590, 595, 615, 616, 637, 641

994

abietum 553, 568 507, 596 508, 530, 585, 597 americana 504, 505, 516, 590 angarensis 508, 531 508, 532 507 aterrima 610 530, 580 53 brachyura 533, 533 553, 568 bosnica 585, 598 brachyura 532 598 chrysospila 611 508 domestica 502, 596 fagorum 596 507 503, 505, 506, 590, 604, 605, 607,609,610,610,611 505, 506, 506, 507, 585, 586, 587, 589, 593, 596, 597, 598, 602 fusco-flavescens 507 gwatkinsii 505, 611 508, 513, 518, 523, 528, 530, 531, 532, hamgyensis 508 533, 533 508, 531 Martinae 505, 541 indochinensis 6 Megalotis 268 intermedia 597 Melodon 817 508, 532 Melanarctos 404 531 404, 408 kamtschadalica 533 Meledes 816 kamtchatica 533 leptorhynchus koreana 6 Meles 493, 494, 498, 500, 500, 803, 816, koreensis 533 817, 5/S, 818 kozlovi 598 aberrans819, 833 kuatunensis 611 altaicus819, 833 latinorum 569 amurensis 817, 819, 836, 837 leucolachnea 597 anakuma 819, 836, 837 553, 569 817, 837 martes 505, 506, 507, 517, 553, 553, 818, 819, 834, 835 atavus 818 554, 557, 562, 563, 567, 568, 569, 569 maculata 507 819,837 mediterranea 598 canescens 819, 833, 836, 837 melampus 505, 533 caucasicus 819, 832 danicus 837
alba
peninsularis
altaica

arsenjevi
asiatica

averini

baicalensis

borealis

bunites

coreensis

ferruginea

flavigula

foina

598 587, 597 596 obscura 508, 532 ochracea 507 ognevi 585, 597 pennanti 503, 505, 516, 590 611 princeps 531 rosanovi 585, 587, 596 ruthena 553, 586 saba 611 sabaneevi 553, 568 508, 532 532 508, 531 568 568 598 szetchuensis 611 tomensis 508, 530 toufoeus 598 szetchuensis 611 533 tungussensis 508, 531 553, 569 508, 532 553 yeniseensis 508 yuenshanensis 611 zibelUna 501, 505, 506, 506, 507, 507,
milleri

nehringi
notialis

sahalinensis

schantaricus

sajanensis

sylvatica

sylvestris

syriaca

tsuensis

uralensis

vitimensis

vulgaris

ilimpiensis

jacutensis

jeniseensis

cavifrons

81

817,

lorenzi

arcalus

arenarius

blanfordi

28

1545
Meles europaeus 819
81
heptneri 819,

gennevauxi 832 leptorhynchus 819, 830, 833, 837 leucurus 817, 837 marianensis 837 melanogenys 819, 837 meles 493, 817, 818, 818, 819, 819, 820,
823, 826, 830, 832, 833, 833, 834, 835,

Mesaxonia 10 Mesonychidae 54
Metatheria 10

Miacidae 54

Miacoidea 54
michnoi
cf.

Mustek

Microaonyx 853 microdon cf. Paraonyx

middendorffii
cf.

Ursus

minor 819, 833 ponticus 837 819 rhodius 837 819, 836 severzovi 819, 833, 833, 835 819, 833 835, 836 819,832 taxus 819 818 tianschanensis 819, 835, 836 819
819,
raddei
schrenkii
sibiricus

835, 836, 836. 837

macrotis

cf.

Lycalopex

minor

cf.

Canis

minuta

cf.

Mustela Mustela

Miosireniae 23

mongolica

cf.

Monotremata 10 montanus cf. Vulpes

monticola
moreotica
cf.
cf.

Lutra, Vulpes

talassicus 819,

Canis

tauricus

mosanensis
mupinensis

cf.

Mustela
Ursus

taxipater 8

cf.

Mustela 493, 498, 501, 505, 639, 636, 636,

641, 643, 848, 690, 691, 704, 705, 723,

thordi

typicus
cf.

meles

Meles

Melinae 498, 802, 803, 816, 851, 854, Melogale498, 817

melli
cf.

Martes

Mellivora 493, 498, 499, 500, 801, 801,

abyssinica 812 broekmani 812 Mellivora buchanani 812 capensis 804, 804, 805, 807, 809, 812, 812 consica 812 812 812 leuconota 812 makwelli 812 pumilio 812 804 812 812 vemayi 812 804, 812
81
cottoni
inaurita

802, 802, 803

indica

ratel

signata

sigulata

wilsoni

Mellivorinae 498, 803

Meiursus 50, 399, 403

ursinus 403,

483

Mephitinae 498, 803

Mephitis 493, 498
meridionalis
cf.

Lutra, Ursus

663, 673 admirata 774 677 alba 585 albica719, 731 671, 676 657 696 691 641, 643, 648, 686, 686, 689, 995 690, 692, 694, 695, 696, 697, 697 amurensis 508, 773 749 anguinae 676 675, 675, 676 712 507 697 aterrima 604 aurea 771 aureola 749 aureus 748, 770 710 508 balkahca 664, 673 bangsi 676 664, 675 biedermanni 732 binominata 719, 730 664, 674, 695
aestiva

724,781,783,787,929

alascensis

algirica

alleghaniensis

alpina

alpinus
altaica

angliae

arctica

asaii

asiatica

astutus

australis

baicalensis

baturini

birulai

1546
Mustela boccamela 644, 650, 654
borealis

553, 604, 719, 730 brachyura 507 buclina719, 731 caledoniae 749 campestris 657 canigula 712 caparioccoi 697 caucasica 655, 730 caucasicus 719 celenda 677 charbinensis 701, 712 cicognanii 679 coreana 712 cylipena718, 731 davidiana 712 dauricus 772 digna 664, 675 655 dombrowskii 644 domestica 585 erminea 636, 637, 641, 642, 648, 663,
dinniki

706, 711 676 kamtschatica 508, 652, 675 663, 675 kanei 675 644, 652 664, 675 641, 697 653 508 770, 773 leucolachnea 585 754, 772 686, 697 854 637, 641, 643, 691, 718, 718,
itatsi700,

kadiacensis

kaneii

karaftensis

karaginensis
katian

690,

kerulenica 644,

koreensis
larvata

lineiventer

longstaffj
lutra

lutreola

719, 720, 722, 726, 728, 729, 730, 731,

663, 664, 665, 666, 669, 671, 672, 673,

eskimo 657 eversmanni 638, 640, 641, 642, 705,

719, 742, 743, 744, 753, 753, 754, 755,
758, 760, 761, 763, 768, 111, 111, 113,

674, 674, 675, 676, 676, 677, 737

fagorum 585 677 ferghanae 674, 676 502, 533, 604, 719 749 585, 587 712 640, 641 736, 743, 749 644, 654 657 719,731 guIo614, 618 gulpsa 677 heptapotamica 771 644, 654, 659 herminea 663 hibemica 676 hodgsoni 712 hungarica 719, 731, 754, 771 657 677 intennedia 585, 587 677
fallenda
flavigula

774,781,929

foetidus

foina

fontanieri

frenata

furo 56,

gale

galinthias

glogeri

882, 884 lymani 667, 674, 676 macrodon 932 manchurica 700, 710, 711, 712 martes 553, 585 martinoi 664, 674 meiampus 507 michnoi 772 miles 700, 710 minima 676 minor 729 minuta 650 mongolica 677 mosanensis 644, 657 mosquensis 737, 749 moupinensis 712 murica 676 namiyei 644, 652, 657 naumovi 664, 675 nehringi 585 641, 761, 769, 770, 774 644, 655 nippon 663, 676 636, 637, 640, 641, 642, 643,
lutris

731,732,924,925

nigripes

nikolskii

nivalis 51,

643, 644, 645, 647, 649, 650, 651, 652, 652, 653, 653, 654, 655, 655, 656, 656, 657, 662, 719

heptneri

iberica

initis

invicta

771 novikovi719, 730, 713 nudipes 641, 690 numidica 657, 671 771 ognevi 664, 674, 737, 749 olympica 677 663, 675, 737, 749
nobilis

occidentalis

orientalis

1547
Mustela pallida 656, 771
polaris

peregusna 781, 783 676 princeps 507, 531 punctata 644, 653 501, 636, 641, 642, 719,
putorius 56,

wyborgensis 718, 729 yesoidsuno 644 502, 507, 508

zibellina 56,

749

MusteUdae 54, 55, 57, 58, 491, 496, 497, 500,785,853,911,924

Mustelinae 497, 498, 502, 637, 641, 801,
803, 817

721, 724, 736, 737, 738, 740, 743, 744,

996

sho712 637, 641, 643, 675. 690, 700,

sibirica

929 pygmaea 51, 644, 652, 653, 656 quelpartis 712 raddei691,696 richardsonii 676 ricinae 676 rixosa 641, 644, 650, 657 robusta 754 rothschildi 737, 749 rupestris 507 russeliana 657 sacana 695, 698 salva 677 sarmatica 781, 783 schnitnikovi 664, 676 seclusa 677 semplei 676
748, 749, 749, 754, 760, 761,

Mutica 10

Mydaus 817
Mylarctos 404

My opus

schisticolor 521

eversmanni 404, 408

Mystacoceti 9

Myrmarctos 404

N
nair cf. Lutra

namijei

cf.

Mustela

Nandinia 49

Nasua908, 911
Nasuella911
Nectogale 497

negans

cf.

Vormela
25
Vulpes

700, 701, 701, 703, 706, 706, 709, 710,

772 676 749 644, 657 677 638, 641 subpalmata 657 553 553 772 924 teberdina 664, 673 temon 686, 697 774 674 tonkinensis 657 663, 675 719, 731 644, 655 730 772 719, 731 vison 641, 643, 723, 724, 924, 925, 928, 930, 931, 932 636, 637, 644, 650, 655, 656,
sibiricus
stabilis

710, 711, 711, 712, 724, 754, 837

Nepus 25
stelleri
cf.

niger

Canis
cf.

nigricans
nivalis cf.

Mustela

stantschinskii

Nyctereutes 60, 67, 63, 66, 67, 68, 68, 69,

stolizkana

streatori

strigidrosa

sylvatica

sylvestris
talassica
tatarica

amurensis 75 75 75 75 procyonoides 75 75 75
69,

911

kalininensis 69,

koreensis

Orestes

59, 63, 64, 69, 69, 70, 71,

74, 75,

ussuriensis

viverrinus 66,

tiarata

tobolica

transbaicalica

obscura

cf.

Vormela
Martes, Mustela, Putorius

transsylvanica
trettaui

Odontoceti 9

ognevi

cf.

turovi

orientalis cf. Canis,

Mustela

tuvinicus

oraata

cf.

Vormela

varina

Otariidae 53

56,

928,

megalotis 63

Otocyon 50, 58, 60, 62, 63

Otocyoninae 62

vulgaris

1548
Oxyaenidae 54

procyonoides

cf.

Nyctereutes

Oxygous 96

Prorastomidae 23
Protosirenidae 23

Proteles51
Prototheria 10

Paenungulata

10, 15, 23,

24

Protungulata 10

Paleomeles 817
pallidior cf.

pruinosus

cf.

Ursus

Vormela
Fennecus
cf.

pallidus cf.

pallipes cf. Canis

pambasileus
pamirensis
Paralutra

Canis

Pusa 882

Pteronura 853, 854, 883

brasiliensis 494,

853

orientalis

884

cf.

Ursus

pusilla cf. Vulpes

854

Putorius 636, 640, 641, 705, 723, 724, 736,

Parameles 817

microdon 854 854

philippsi

Paraonyx 498, 853, 854, 883 congica 853

Parataxidea 817, 818

Paraxonia 10

peregusna

cf.

Vormela
Martes

Perissodactyla 10, 11, 54

peninsularis
philippsi cf.
cf.

Paraonyx

Pholidota 10

Pinnipedia 10, 11,53,54

Pteronura 498
piriformis
piscator
cf.

Putorius

cf.

Ursus

planicola

cf.

Vulpes

Plesiogale

640

brachygnatus 616 monspessulanum 616

Plesiomeles 817

Plesiogulo616

Poecilictis 498, 783,

libica

785

783

Poecilogale 498, 783, 785

Polyprotodontia 9

Potamotherium 854, 883

Potos907, 908, 911
Prosimiae 9
Primates 10

Proboscidea 10, 23, 24, 54

Procynodictis 60

997

cancrivorus 912, 914, 919 56, 912, 913, 913, 915, 916, 977,
lotor

Procyon 69, 908, 970, 911, 912, 912

919
Procyonidae 54, 55, 57, 60, 69, 402, 497,

907,909,911 Procyoninae911,912

686 amurensis 754 663 686 aureus 754, 761,772 caucasicus 644 dauricus 754 644 erminea 663 eversmanni 753, 754 ferghanae 663 736 700 743 heptapotamicus 754 kamtschatica 663 663 kanei 663 753, 770 michnoi 753 644 754 754 ognevi 737 737 644, 754 737 736, 737, 739, 754, 761, 767, 772 pygmaeus 644 754 663 754 736, 739, 767 754 754 verus 736 736
alpinus
arcticus

742, 783, 929

astutus

dinniki

foetidus

fontanieri

furo

kaneii

larvatus

nivalis

nobilis

occidentalis

orientalis

pallidus

piriformis

putorius

satunini

sibirica

sibiricus

stantschinskii

talassicus

tuvinicus

vulgaris

putorius

cf.

Mustela

1549
pygmaea
Mustela

cf.

R
rixosa
cf.

talassica cf.

Mustela
521

Tamias
Mustela
10, 11

sibiricus

tatarica cf.

Mustela

Rodentia

Taxidea 493, 494, 498, 817, 818, 819

Rohrwolf 140
rothschildi cf. Mustela

Taxodon 817 Taxus 816

Tayra barbara 616
teberdina
cf.

rufus

cf.

Canis

riippeli cf.

Fennecus

Mustela

Rytina 25

temon

cf.

Mustela

cetacea 25

Rytina borealis 25

Thalarctos 403, 404, 406, 455

Rytiodinae 23

404 Thalassarctos 404 jenaensis 455 455

polaris

spitzbergenis

Thalassiarchus 404

Thos 96, 99

aureus 96
sacana
cf.

Sacalius 96

vulgaris
cf.

96
cf.

thous

Lycalopex

Mustela

tianschanensis
tiarata cf.

Meles

SchaefFia 99
seistanica cf. Lutra

Mustela
cf.

tuvinicus

Mustela

ussuricus 478
severzovi
cf.

Selenarctos 403, 404, 406, 414,

478

tobolica

cf.

Mustela

Tomarctos 268
Theria 10

Meles

venaticus 63
sibirica cf.

Speothos 62

998 Tarsioidea 9
transsylvanica
cf.

Mustela

Mustela
Ursus
Mellivora

Tremarctos 399, 403, 404, 406

sibiricus cf. Gulo, Meles,

ornatus 40], 403, 406

trettaui cf.

signata

cf.

Mustela

simensis 62
Simiae 9
Sirenia 10,

Simenia 62, 68, 99

Trichechidae 15, 17, 23, 24

Trichechiformes 23, 25

Trichechus 15, 17, 18, 23, 24

Simocyoniae 62, 63, 383

11,15,27,22,54

seistanica cf. Lutra

Sivaonyx 854

25 inuguis manatus senegalensis

borealis
tschiliensis cf.

16, 20,

24
25

20, 24,

20,

24

Smilodon 50
Speothoninae 62

Canis

Tubulidentata 10, 54

venaticus

Speothos 58, 60, 62

59,

turcmenicus

cf.

Vulpes

63
cf.

tundrarum

cf.

Canis

Spilogale 493, 498

spitzbergenensis

turovi cf. Mustela

Alopex

Stelleras 25

borealis 25

Stenoplesictinae 497

Stromeriella 817

ungava

cf.

Alopex
cf.

Subungulata 22
syriaca
cf.

Unguiculata 10
ungavensis

Vormela
Ursus
cf.

Ursus

syriacus

cf.

Ursarctos 404

szetchuensis

Martes

Urocyon

59,

62

1550
Urocyon cinereo-argentatus 63
Ursavus 402
Ursi campestres

419

Ursidae 54, 55, 56, 57, 60, 397, 401, 402,

911

albus618 americanus 399, 400, 404, 405, 406 399, 404, 406, 407, 407, 408, 411,
arctos

Ursus 398, 401, 403, 404, 404, 406, 407

412, 415, 422, 424, 428, 429, 430, 431,

431, 432, 433, 433, 434, 435, 436, 436,

aureus 407 408, 429 beringiana 407, 436 beringianus 435, 436 brunneus 407, 436 cadaverinus 407 caucasicus 408, 431 436 collaris407, 429, 431 crowtheri 428, 437 43 eogroenlandicus 467 formicarius 407 formosanus 485 gedrosianus 485 grandis 408 407 groenlandicus 469 gulo615, 618 gyas 428, 436 hypernefes 408 804 407, 434, 435, 439, 453 japonicus 485 jenaensis 467 406 kamtschatica 407 kolymensis 408, 429 469 lagomyarius 408 485 404, 408, 436, 436 408, 433 leuconyx 408, 434 407 lotor912, 913 618 mandchuricus 408 marinus 455, 460, 467, 468 maritimus 404, 406, 407, 455, 455,
baicalensis

437, 439, 451, 453, 459

cavifrons

dinniki

meles816, 819 485 407, 431, 431, 432, 433, 434, 459 minor 407 428, 436 mupinensis 485 myrmecophagus 407 407 normalis 407, 408 omatus 404 pamirensis 408, 434, 434 408 407, 435, 436, 437, 459 pruinosus 404, 407, 434, 435, 437 455, 467 408 scandinavicus 408 408 smirnovi 408, 431 467 408 407, 431, 432, 433, 434, 435, 459 404, 411, 478, 479, 481, 483,
melli

meridionalis

middendorffii

niger

persicus
piscator

polaris

rossicus

sibiricus

spitzbergenensis

stenorostis

syriacus

thibetanus

484, 485, 485

Ursus tibetanus 399, 403, 404, 406, 407,

griseus

indicus

isabellinus

478 tyrannus 460 ungavensis 469 485 408, 429, 432, 433, 436, 437, 459 yessoensis 404, 408
torquatus
ussuricus

414, 437

yeniseensis

ussuricus

cf.

Ursus

ussuriensis cf. Nyctereutes

jessoensis

labradorensis

vancouverensis
Vetularctos

cf.

Gulo

laniger

406 Vishnuonyx 854

lasiotus

lasistanicus

iongirostris

capensisSOl, 804 Iutreola636, 718 ratel801

Viverridae 54, 55, 497
viverrinus
cf.

Viverra aterrima 884

luscus

Nyctereutes

411,

456, 458, 461, 465, 467. 468

alpherakyi 794 798

chinensis

Vormela 498. 501, 501, 781, 781, 782, 783

1551
Vormela euxina 784, 794
-

999

intermedia 794 koshevnikovi 783, 794, 196, 797 negans 784, 796, 797 obscura 784, 794 omata 784 796 784, peregusna 783, 783, 784, 784, 785, 796, 797, 798 794, sarmatica 784, 794 784, 797 tedschenica 783, 794
795,
pallidior

flavescens 66, 319, 333, 334, 337, 340,

340, 360
-

fulva 327, 341

griffithi

341

harrimani 341

-hoole 341
-

795,

huli

320

752,

induta 341

788, 790, 793,

795,

ischnusae 341
jacutensis 320,

339

syriaca

-japonica 341
-

Vulgaris

cf.

Mustela

Vulpes 62, 63, 68, 69, 99, 194, 196, 265,

abietorum 341 aegyptiaca 341 341 alopex 319 alpherakyi 319,338, 338 338 anadyrensis 319, 339 341 341 341 bangsi 341 barbara 341 bengalensis 327 beringiana 339, 339, 341 beringianus 196 cana 266, 268, 269, 269, 270, 271, 272, 327 caragan 339, 340 cascadensis 341 caucasica 338 chama 268 340 communis 265 319 corsac 273, 276, 279, 280, 327 341 320, crymea-montana 338 crymensis 319 339, 339 341 337, 339, 340 339 325, 281 340 320, 272, 327 femlatus 63,281
alascenis
alticola 320,

265, 268,

277

kalmykorum 273, 280 kamtschadensis 319, 339 kamtschatica 319, 339 karagan 320
kenaiensis 341

kiyomassai 320, 339

krymea-montana 319
krymensis 338
iagopus 196
bengalensis 63

-kurdistanica319, 338
-

anatolica

leucopus 319

arabica

macroura 341
melanotus 339

atiantica

montana 341 monticola 341

necator 341
nigra 273, 281

268, 272,

nigricans

269

63, 66,

269,
-

ochroxantha 320, 340

paiestinae 341

336,

pamirensis 320, 340

peculiosa 341

3 1 9,

planicola 341
pusilla 341

cinerascens 319,

pusillus319
regalis 341

coloregriseo

rabricosa 341
riippeli

63, 66, 266, 268,

274, 275,

268

281, 315, 337,

schrenkii 320, 339,

341 336

crucigera 319,

scorodumovi 273, 281

septentrionalis 319,

daurica 320,

339 sibiricus 319

sibirica

deletrix

silacea 341

diluta 320,

337,

splendens319, 340, 340

splendidissima 320, 339, 341
stepensis

dolichocrania 320,
eckloni

336,

319,337,338
340

ferganensis

333,

tarimensis 319
tobolica 320, 339.
tschiliensis

ferrilata 268,

319

1552
Vulpes turcmenica 273, 280, 315
ussuriensis

319, 339 velox 63 319 vulpes

vulgaris

yeniseensis

cf.

Martes, Ursus
cf.

yuenschanensis

Martes

56, 58, 63, 64, 66, 66, 266, 268,

318, 319, 320, 32J, 326, 327, 329, 330,

333, 334, 336, 336, 337, 337, 337, 338,

339,339,340,340.341,560 Vuipicanis 96

zerdus

cf.

Fennecus
Martes

zibellina cf.

W
wilsoni
cf.

Zibellina

502 783

Zorilla libyca

Mellivora

Other Books of Interest

MAMMALS OF THE SOVIET UNION

Volume
II,

Part 2
is

monograph, and is a continuation, which was devoted to sea cows and carnivores. It contains species descriptions of terrestrial carnivores and detailed information on their external morphology, skull, body measurements and
This volume,
of Volume
II,

part of a three-volume
1,

Part

other data, affinities with other species, geographic distribution in the

historic past

and today, geographic

variation, practical significance,

biology. Descriptions are presented for orders, families,

and and genera, and

keys given for their identification.

The book
illustrations

is

richly illustrated with photographs, sketches,

painter, A.N.

by the famous wild life zoologist-artist, N.N. Kondakov.

and colored Komarov, and the

The
work
is

results of original scientific studies are published here for the first

time, providing readers a vast wealth of material heretofore

unknown. This

intended for teachers and students of faculties of biology and geography in universities, as well as pedagogic, agricultural, and forest
institutes, similar organizations,

workers engaged in game,

study.

fur, forest
all

and

fish trades; those interested in the conservation of nature,

and

persons

interested in zoology

and nature

MAMMALS OF THE SOVIET UNION

Volume
II,

Part 3
is

This volume

fourth book of Mammals of the Soviet Union, representing the

it is

third part of the second volume;

devoted to descriptions of the orders

part,

of Soviet aquatic

mammals

pinnipeds (Pinnipedia) and, in

i.e.,

cetaceans (Cetacea), toothed whales (Odontoceti). In the sequence of

descriptions from the "higher" to the "lower" orders adopted in this series,

pinnipeds should have preceded carnivores,

the second book.

should have appeared in

The grouping of the orders at a higher level is given after G.G. Simpson (1945). The sequence of genera and species within the orders
has been retained as before,
i.e.,

from the

less specialized to

the

more

specialized.

The order of pinnipeds or seals (Pinnipedia) is described in this volume. The cohort of ungulates and carnivores represented in soviet fauna by
orders of artiodactyls, and perissodactyls, sirenians, carnivores, and

pinnipeds thus came to an end and the cohort of whales (Mutica) commences. The toothed whales are described in this volume.

While

it

has not always been possible to maintain a totally uniform

description of the genera and species as in the volumes already published,

in spite of every effort to do so, the sequence has been adhered to, with

some exceptions, in the case of Pinnipedia. But, it was impossible to use same format in describing the toothed whales (Odontoceti). Only a brief morphological description has been given for many species, which is more or less adequate for identifying the species. General information on their distribution and fragmentary biological data are also given. In some cases
the total absence of such information
is

indicated.