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Full text of "Mammals of the Soviet Union"

Mammals 

of the ^ 

Soviet Union 



VOLUME П 

^ Rut lb 



VG. Heptner 



N. E Naumov 



т.- '^- 




^-s^-- 



The present book, of the series Mammals of 
the Soviet Union: Sirenia and Carnivora, is the 
fourth volume to be published in English. 
Previously Volume I — Artiodactyla and 
Perissodactyla, Volume II, Part 2 — Carnivora 
(Hyenas and Cats) and Volume II, Part 3 — 
Pinnipeds and Toothed Whales were published 
in 1988, 1992, 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. The 
first (Part la) consisted of the Steller's sea 
cow, the now-extinct marine 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 
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 point: since the appearance of 
Volume II, Part 2, the Soviet Union has 
disappeared. However, the series was written 
in the context of the former "union of 
republics," and it would be confusing to re- 
edit the text to conform to the current 
political reality. 



Mammals of the Soviet Union 

Volume II 
Part lb 



MAMMALS OF THE SOVIET UNION 

In Three 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: v. 2, pt. la, p. 

Vol. 2, pt. 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) 

vol. 2. — pt. lb. Carnivora (Weasels; Additional species) 

vol. 2. — pt. 2. Carnivora (Hyaenas and cats). 

1. Mammals — Soviet Union. I. Nasimovich, A. A. II. Bannikov, 
AndreT Grigor'evich. Ill Hoffmann, Robert S. IV Title. 
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.) 735 
Key for Identification of Genera in the 

Family Mustelidae 746 

Subfamily Mustelinae 749 

Genus Martes Pinel, 1792 (Martens) 749 
Key for Identification of Species in the 

Genus Martes 755 

Martes {Martes) zibellina (Linnaeus, 1758) 

(Sable) 757 

Diagnosis 759 

Description 759 

Systematic Position 770 

Geographic Distribution 770 

Geographic Variation 787 

Biology 797 

' ' Practical Significance 823 

Martes {Martes) martes (Linnaeus, 1758) 

(Forest Marten) 825 

Diagnosis 826 

Description 826 

Systematic Position 836 

Geographic Distribution 836 

Geographic Variation 847 

Biology 850 

Practical Significance 872 

Martes {Martes) foina Erxleben, 1777 

(Stone Marten) 874 

Diagnosis 875 

Description 876 

Systematic Position 882 



Geographic Distribution 883 

Geographic Variation 891 

Biology 895 

Practical Significance 904 

Martes (Charronia) flavigula Boddaert, 1785 

(Kharza, or Yellow-throated Marten) 905 

Diagnosis 905 

Description 905 

Systematic Position 910 

Geographic Distribution 910 

Geographic Variation 912 

Biology 914 

Practical Significance 919 

Genus Gulo Storr, 1780 920 

Gulo gulo (Linnaeus, 1758) (Wolverine) 925 

Diagnosis 926 

Description 926 

Systematic Position 931 

Geographic Distribution 932 

Geographic Variation 940 

Biology 943 

Practical Significance 953 

Genus Mustela (Linnaeus, 1758) 954 

Key for Identification of Species in the Genus 



Mustela 






962 


Mustela {Mustela) nivalis Linnaeus, 


1766 




(Weasel) 






965 


Diagnosis 






966 


Description 






967 


Systematic Position 






972 


Geographic Distribution 






972 


Geographic Variation 






975 


Biology 






985 


Practical Significance 






995 


Mustela {Mustela) erminea 


Linnaeus 


, 1758 




(Ermine) 






995 


Diagnosis 






997 


Description 






997 


Systematic Position 






1003 


Geographic Distribution 






1003 



Vll 



Geographic Variation 


1008 


Biology 


1016 


Practical Significance 


1029 


Mustela (Mustela) altaica Pallas, 1811 




(Solongoi) 


1030 


Diagnosis 


1031 


Description 


1031 


Systematic Position 


1036 


Geographic Distribution 


1037 


Geographic Variation 


1043 


Biology 


1047 


Practical Significance 


1051 


Mustela {Mustela) sibirica Pallas, 1773 




(Kolonok or Siberian Weasel) 


1051 


Diagnosis 


1052 


Description 


1052 


Systematic Position 


1058 


Geographic Distribution 


1058 


Geographic Variation 


1065 


Biology 


1069 


Practical Significance 


1078 


Mustela {Mustela) lutreola (Linnaeus, 1761) 




(Russian, or European Mink) 


1078 


Diagnosis 


1080 


Description 


1080 


Systematic Position 


1086 


Geographic Distribution 


1088 


Geographic Variation 


1094 


Biology 


1099 


Practical Significance 


1106 


Mustela {Putorius) putorius (Linnaeus, 1758) 




(Forest, or Black, Polecat) 


1107 


Diagnosis 


1108 


Description 


1108 


Systematic Position 


1115 


Geographic Distribution 


1117 


Geographic Variation 


1124 


Biology 


1126 


Practical Significance 


1132 



vin 



Mustela (Putorius) eversmanni Lesson, 1827 




(Steppe, or White, Polecat) 


1133 


Diagnosis 


1134 


Description 


1135 


Systematic Position 


1142 


Geographic Distribution 


1145 


Geographic Variation 


1158 


Biology 


1165 


Practical Significance 


1175 


Genus Vormela Blasius, 1884 


1176 


Vormela peregusna (Giildenstaedt, 1770) 




(Marbled Polecat, Perevyazka) 


1180 


Diagnosis 


1181 


Description 


1181 


Systematic Position 


1188 


Geographic Distribution 


1188 


Geographic Variation 


1194 


Biology 


1202 


Practical Significance 


1206 


Genus Mellivora Storr, 1780 


1207 


Mellivora capensis (Schreber, 1776) 




(Ratel, or Honey Badger) 


1211 


Diagnosis 


1211 


Description 


1211 


Systematic Position 


1217 


Geographic Distribution 


1217 


Geographic Variation 


1222 


Biology 


1223 


Practical Significance 


1228 


Subfamily Melinae 


1228 


Genus Meles Brisson, 1762 [=Boddaert, 1785] 


1228 


Meles meles (Linnaeus, 1758) 




([Eurasian] Badger) 


1232 


Diagnosis 


1234 


Description 


1234 


Systematic Position 


1242 


Geographic Distribution 


1242 


Geographic Variation 


1249 


Biology 


1262 


Practical Significance 


1281 



IX 



Subfamily Lutrinae 




1283 


Genus Lutra Brisson, 1762 [=Brunnich, 1771] 


1283 


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


1288 


Diagnosis 




1289 


Description 




1289 


Geographic Distribution 




1297 


Geographic Variation 




1305 


Biology 




1310 


Practical Significance 




1327 


Genus Enhydra Flemming, 1822 




1330 


Enhydra lutris (Linnaeus, 1758) (Sea 


Otter, 


1333 


or Kalan) 






Diagnosis 




1333 


Description 




1334 


Systematic Position 




1343 


Geographic Distribution 




1343 


Geographic Variation 




1348 


Biology 




1350 


Practical Significance 




1365 


Supplement to the Order of Carnivores 




1366 


Family Procyonidae (Raccoons, etc.) 




1366 


Subfamily Procyoninae Gill, 1872 




1373 


Genus Procyon Storr, 1780 




1373 


Procyon {Procyon) lotor (Linnaeus, 


, 1758) 




(American Raccoon) 




1374 


Diagnosis 




1374 


Description 




1374 


Systematic Position 




1377 


Geographic Distribution 




1377 


Geographic Variation 




1383 


Biology 




1383 


Practical Significance 




1390 


Family Mustelidae 






Mustela (Mustela) vision Schreber, 


1777 




(American Mink) 




1391 


Diagnosis 




1391 


Description 




1392 


Systematic Position 




1397 



*In Russian original, Lutra (Lutra) — Sci. Ed. 



Geographic Distribution 1399 

Geographic Variation 1403 

Biology 1405 

Practical Significance 1415 

LITERATURE CITED 1416 

INDEX OF LATIN NAMES OF TAXA* 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 mod- 
erate 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. 

The limbs are digitigrade or semi-plantigrade, but several are 
plantigrade with five digits on the fore- and hind-limbs. The ventral 
surface of the hind foot and forepaw are covered with hairs (as is, 
usually, the skin of the interdigital 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, it is more proportional, but the facial portion 
is always shorter than the braincase. In small forms, the tubercles, 
crests, etc. of the skull are usually weakly developed, and in large 

493 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 is quite high, with the 
upper profile an arciform line. The mastoidal processes are usually 
small, as are the paroccipitals. The auditory bullae are quite large, 
but flattened or only weakly inflated, usually without a septum. 
The alisphenoid canal is absent. The orbit is large, and the tempo- 
ral fossa is of great size. 



736 



The second upper molar is absent. The complete dental for- 

3 14 1 
mula is I — C-P — M — = 38. This formula is not encountered in 

all species of the family (developed in martens, Maries; wolverine, 
Gulo; badgers s.s., Meles, Helictis, Arctonyx and others). A re- 
duction in tooth number often occurs in the premolars and molars. 



This occurs in various combinations 



3 1 



4 1 
P - M - = 36 

3 2 



(otters, Lutra); 



3 1 



3 1 
P-M — = 34 (polecats, weasels, Mustela, 



Eyra, Galictis; American badgers, Taxidea; skunks Spilogale, 
Mephitis) and even 



ilci 

3 1 



2 1 
P — M — = 32 (skunks, Conepatus) and 



lici 

3 1 



3 1 
P — M - = 32 (ratel, Mellivora). In one case (sea otter, or 



kalan, Enhydra), reduction occurs in the number of incisors re- 

3 13 1 
tained, and the formula takes the form I-C-P-M- = 32. The great- 
est reduction, however, is reached in the number of molars and 

premolars 



3 1 



2 1 
p — 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 — the 

4 3 4 3 
dental formula may be P-P-P-P- = 38 = 36 = 34 (badger, Meles 
•^4433 

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 an os penis, usually thin and long, sometimes strongly 
curved, divided 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 
more or less 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. 

The pelage, in relation to different kinds of biological types 
and the regions of occurrence, is quite variable. Some northern 
494 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 back- 
ground (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 
somewhat larger than females. Age dimorphism in color is absent 
or is expressed only in the first juvenile pelage. Seasonal dimor- 
phism is only manifested in northern forms and may be very sharp, 
with regard to length, density and other fur properties. Fur color is 
somewhat variable. In extreme cases, the 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 numer- 
ous species of the family are extremely diverse, although they may 
be divided into several biological types (see below). A feature 
common to all lies in their relatively short limbs. The majority are 
characterized by a thin 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 character- 
istic. 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 — martens and ilka*; 
excellent swimmers and divers, the truly amphibious fresh-water 
otters with very long and flexible body. Specialization reaches its 
extreme degree in marine-dwelling sea otter living on the sea- 
shores, which possesses pinniped characteristics (structure of the 
hind limbs). 

These forms with long to very long trunk and short limbs are 
unable to move at a trot 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. 
More or less "neutral" are the little specialized forms (South-Asian 
badgers), but the more developed are represented by the relatively 
large animals of 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 rep- 
resentatives of the family. This is connected with their remarkable 
passive defence (exuding a stinking fluid from the anal glands). 
The wolverine is large and massive, but a good climber. 



♦Russian name for North American fisher, M. pennanti — Sci. Ed. 
**Literally, "stinkers" — 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 cm 
(South- American otter, Pteronura brasiliensis) and even 150 cm 
(sea otter). The latter has a weight of up to 40 kg. The smallest 
495 form has a body length of about 15 cm, a weight of about 100 g 
or less (weasels of the group pygmaea* — rixosa). The ratio of the 
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 common, or neighboring, 
burrows. Species leading a sedentary mode of life do not occur in 
the family (data on some American otters are not well defined). 

Trophic specialization is quite variable. There are trends to- 
ward sharply pronounced energetic carnivores feeding on warm- 
blooded vertebrates (chiefly rodents — weasels, polecats, etc.) and 
strongly restricted ichthyophages (otters). With one or another de- 
fined 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 in feeding on sea-urchins (sea otter, or 
kalan). 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. 
Of the sense organs, hearing is, apparently, the most developed, 
and in some, smell also. The general level of psychological [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 level to extreme heights. 

*Spelled pygmea in Russian original — Sci. Ed. 



740 

Their geographical distribution is very extensive — nearly the 
entire world. The range includes the entirety of the South Ameri- 
can continent, Central and all of North America, including the 
whole Arctic archipelago. In the western hemisphere, representa- 
tives of the family are absent only on the Falkland and Galapagos 
islands, 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 
Greenland, the range area occupies only the very northern part of 
the island east of Ross Strait and its eastern coastal zone south- 
ward, somewhat more southerly than latitude 70" N. lat. (below 
Scoresby Sound). In the Old World, the range includes the whole 
of Africa, Europe and the mainland of Asia. Species of the family 
are absent on Iceland and the majority of the Arctic Islands 
(Spitsbergen, Franz Josef Land, Novaya Zemlya, Severnaya Zemlya 
and Wrangel Island), but are encountered in Kolguev, Vaigach and 
the New Siberian archipelago. In the Far East, the range encom- 
passes 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, 
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 quite strongly changed. However, the range of 
the family as a whole has not changed substantially in the last 
century. 

In spite of its species richness and especially its diversity of 
forms, the family represents a well separated group. It is distinctly 
demarcated from other families within the order and in the group 
Canoidea, and 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. Among the viverrids, only one 
genus — Nectogale — is analogous to otters though not more spe- 
cialized. The external similarity of 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 is closely 
related to raccoons (Procyonidae) and is relatively far from the 
bear and wolf families. 

The family itself represents a quite ancient group — its first 
representatives were found in the early Oligocene together with 
the most primitive representatives of the wolf, viverrid and cat 
families. They exhibit the greatest closeness to primitive forms of 
the wolf family. However, differentiation of the mentioned fami- 
lies 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 
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. Differen- 
tiation 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 
Mustelidae were known from the lower Oligocene, in Africa they 
are found from the middle Pliocene, and in South America — only 
in the Pleistocene. 

Concerning internal structure, the contemporary family is ex- 
tremely varied, and until now, there has been no accepted and 
really satisfactory division of it 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 
diversity of forms, the diversity of characters and complexity of 
radiation in the family. The view has even been expressed that the 
present Mustelidae does not comprise a phylogenetically united 
group, but is of polyphyletic nature. The heterogeneous character 
of the group becomes even clearer if fossil forms are taken into 
account; if the extreme subdivision of existing forms (15 sub- 
families) is followed, not less than 15 extinct subfamilies must 
also be accepted (Simpson, 1945). 

It is most probable, that the most natural division of the family 
will turn out to be into 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 
subfamilies — the extinct Leptarctinae and 5 contemporary ones. It 
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). 
498 The subfamily of true weasels, Mustelinae, includes the fol- 
lowing 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 
species); Eira (tayra — North and South America; 1 species); Galictis 
(grison, including Grisonella — North and South America; 2 spe- 
cies); Lyncodon (South America; 1 species); Ictonyx (African pole- 
cat — Africa; 2-3 species); Poecilictis (Africa; 1 species); 
Poecilogale (Africa; 1 species); Gulo (wolverine — Eurasia and 
North America; 1 species). 

The subfamily of honey-badgers, Mellivorinae, includes one 
genus Mellivora (Africa, Asia; 1 species). 

The subfamily of badgers, Melinae, comprises the following 
genera: Meles (Old World badgers — Eurasia; 1 species); Arctonyx 
(hog badger — eastern Tibet and southeastern Asia; 1 species); 

'The following list of genera is not in accord with data of Simpson (1945). His 
revision based on various data led to reduction in the number of genera, mainly by 
transferring some to the rank of subgenus. It is apparently possible that further reduc- 
tion will take place. Genus Grammogale Cabrera (Cabrera, 1940, 1957), not recog- 
nized by Simpson, is here set aside. The number of species established from different 
sources, cannot be considered precise. Actually, they are probably somewhat fewer. 



744 

Taxidea (American badger — North America; 1 species); Melo- 
gale, including Helictis ("polecat" [ferret] badgers — southern Asia; 
4 species). 

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 otters, Lutrinae, includes the following gen- 
era: Lutra (including Lutrogale, true otters — Eurasia, African, North 
and South America; 10 species); Pteronura (giant otters — South 
America; 1 species); Aonyx (including Amblonyx; "clawless" 
otters — Africa, Asia; 2 species); Paraonyx (Africa; 3 species); 
Enhydra (sea otter, or kalan — northern Pacific Ocean; 1 species). 
In the entire family, therefore, there are 24 genera and about 70 
species. This constitutes about 1/3 of the species of the order. It is 
more probable that there are about 65 species. As regards the 
number of genera and species, the family is 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); 
six are Asian and Eurasian, three — North American, two — South 
American (Grammogale is not taken into account); 2 — Afro-Asian, 
3 — North and South American; 1 — distributed in Eurasia, North 
and South America; and 1 — in Eurasia, the Americas and Africa. 
Therefore, as regards the number of genera, the fauna of the Old 
World and especially of Asia is the richest and most heterogene- 
ous. 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 — Eurasia (about 17 
endemic species and several others common to the America and 
Africa). It is remarkable, that in spite of only two endemic genera, 
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 practical significance of the family is very great. It is one 
of the most important, if not the most important, group of carni- 
vores. 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 of them have even played a great historical role — the open- 
499 ing up of Russian Siberia was first of all connected with sable, 
whereas the northern Far East and the shore of the northern part 
of the Pacific Ocean, including the American — with sea otter. The 
penetration of Europeans into northern North America from the 
south, was also determined chiefly by furs, among which an im- 
portant 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 
the main fur species in tropical and equatorial countries which, 
owing to their natural conditions, are very poor in fur-bearing 
animals. There are some less valuable fur-bearing species (badg- 
ers, etc.) which, in total, also play a certain role. Finally, the very 
great, and at the present time, the main product of fur farming 
consists of American mink raised in various, artificially obtained 
color forms (mutations). The mink is now a more important fur- 
producing animal than the silver-black fox. Sable, martens, and skunks 
are also raised in captivity, but their importance is very small. 

Many species specializing in feeding on small rodents (wea- 
sels, 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 known 
role in their capacity to exterminate fish and cause harm to fish- 
farming. Finally, a few species serve as objects of sport hunting. 
The fat of some is used in folk medicine. 

In the fauna of the USSR are represented 4 out of 5 sub- 
families (80%), 8 of 24 genera (33%) and 17 species (not counting 
acclimatized) of 70 (about 23%) (V.H.). 

*Meaning unclear — Sci. Ed. 



746 

Key for Identification of Genera of the Family Mustelidae 

1 (2). Digits of the forelimb conjoined, and entire paw repre- 
sents a single unit. Hind limbs transformed into elon- 
gated wide flippers, in which digit V is longest, and evenly 
covered with fur dorsally and ventrally. In each half of 
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 otter or kalan, Enhydra (page 1330). 

2(1). 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 tu- 
bercles or with sharp cusps. 

3 (4). External ear conch absent or rudimentary; entire lower 
body, limbs, side of the head and muzzle black, upper 
side of body is grayish-white, color boundaries of both 
fields 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 4 (3). External ear conch present, color of one sort or another. 
Molars and premolars 5-6 in lower jaw. 

5 (6). Ventral surface of body black or blackish-brown, dorsal 

light, dirty-gray (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, if ventral color black or darker than dor- 

sal, then back has a clearly developed yellow or yellow- 
ish tone. Posterior upper molar not larger than preceding 
one. 

7 (8). Digits united by broad membrane which is nearly com- 

pletely devoid of hairs, 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 
cm. Last molar tooth approximately equal in dimensions 

^Crown viewed from above. 



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 — Genus of badger, Meles; 3 — Genus of 
otters, Lutra; 4 — Genus of wolverine, Gulo. 

to preceding one. First upper premolar pushed away from 
toothrow and lies at inner side of canine (Figure 186) 
genus of otters, Lutra (page 1283). 

8 (7). Swimming membranes between digits absent, or are very 

weakly developed, covered with hairs and do not reach 
end of 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 mm. Tail 

relatively short (about 20 cm). Coloration evenly brown 

501 with light bracket-form bands of different intensity 



*Misspelled in Russian original — Sci. Ed. 



748 




Fig. 186. Position of canine and first premolar of the upper jaw of representative of 
genus of otters, Lutra. Sketch by N.N. Kondakov. 

passing along sides and across croup from shoulder to 
shoulder genus of wolverines, Gulo (page 920). 

10(9). Dimensions smaller — body length less than 65 cm, 
condylobasal length of skull less than 1 15 mm. Tail length 
variable; coloration otherwise. 

11 (12). In upper jaw behind canine 5 teeth (4 premolars and 1 
molar), in lower — 6 (4 premolars, 2 molars). On posterior 
side of main cusp of last premolar (third tooth from rear) 
the lower 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 — yellow with irregular dark spots 
and stripes, 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 




502 



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. 

Subfamily of Weasels 

Subfamilia Mustelinae Gill, 1872 

Genus of Martens 



Genus Martes Pinel, 1792 

\115. Martes. Frisch. Natur-Syst. d. vierfuss. Thiere, p. 11. 

According to decision of the International Commission on 

Nomenclature, names in this edition have no nomenclatural 

significance. 
1792. Martes. Pinel. Actes Soc. Hist. Nat. Paris, 1, p. 55. Martes 

domestica Pinel = Martes foina Erxleben. 
1829. Zibellina. Каир. Entw. Gesch. u. Natur. Syst. Europ. Thierw., 

1, p.p. 31, 34. Mustela zibellina Linnaeus. 
1865. Charronia. Gray. Proc. Zool. Soc. London, p. 108. Mustela 

flavigula Boddaert. 
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 rela- 
tively elongated, zygomatic arches weak, not strongly diverging, 
laterally interorbital and postorbital constrictions weakly defined. 



750 

Supraorbital processes small. Upper profile of skull gently sloping, 
slightly convex. Convexity of fronto-nasal region at level of 
infraorbital foramena weakly developed. Diameter of infraorbital 
foramena almost equal to 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. 

3 14 1 
Dental formula 1-СуР-М- = 38. 

First premolars very small and sometimes shed, but an alveo- 
lus or its traces remains. On main apex of last lower premolar 
(anterior to carnassial tooth — third tooth posteriorly), there is a 
small additional cusp on the inner side. On inner side of median 
apex of 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 moder- 
ately short — general body structure fairly slender and relatively 
proportional. Tail quite long — in various species from 1/3 to 2/3 of 
body length. Head comparatively large with large protruding ears, 
broad at base, 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 — 
fore- 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 combina- 
tion of white, black, yellow and brown. On the throat and chest, 
there is usually a lighter area. 
503 Seasonal dimorphism in character of fur very pronounced; in 

winter, soles of the fore- and hind feet are entirely or almost en- 
tirely covered with dense hairs. Seasonal variations in color slight. 
Sexual differences in color are absent; but dimensions of males 
average larger than females, and in several forms are quite consid- 
erable; 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 monogamous. Litters from 1-2 and 
3-4, up to 8. Shelters are mainly in hollows and in tree trunks, but 
also in rock clefts. They are solitary animals. They are predators, 
feeding on flesh, chiefly of small rodents, but sometimes also on 
larger warm-blooded animals, even small ungulates (yellow-throated 
marten). 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. 

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 cm and a weight up to 6 kg. 

The range of the genus is vast and covers a considerable part 
of the temperate and cold-temperate zones of the northern hemi- 
sphere, 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 for- 
ests [southward] to Spain, Italy, Greece, Crete, Asia Minor, Syria, 
Palestine, Iran, West Pakistan, Kashmir, Punjab, Himalayas, Indo- 
china, the Moluccas, Sumatra, Banka [Island], Java and Kalimantan 
(Borneo). A small part of the range, in southern India, is cut off 
from the general area of habitation (for details, see below under 
descriptions of individual species). 

In North America, the range in the north is limited by the 
northern border of forest vegetation. The southern border is repre- 
sented by a meandering line, extending from the Atlantic coast 
westwards through the states of Connecticut, New Jersey, Vir- 
ginia, northern North 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- 
ish Colombia and from there and from Alberta, the range gives off 
two large extensions southward along montane regions. One ex- 
tends along the coastal regions — along the western parts of the 
states of Washington, Oregon and to the central parts of Califor- 
nia, 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 
the northern parts of New Mexico. In the range is included New- 
foundland, 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 sketch- 
ing the boundaries in the steppe and desert regions of Asia, con- 
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- 
spond everywhere to the restored range cited. 
505 Within the limits of the family, the genus itself constitutes a 

well-isolated group, characterized by a series of features, most of 
all, by the dental formula. Craniologically, the genus is entirely uni- 
form 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 
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 now rejected. 

Many authors divided the genus into two — the 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 is more correct to separate it only as subgenus. There are no 
essential craniological differences in the yellow-throated marten. 





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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 its radiation, is quite 
difficult. 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 be- 
tween 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- 
lished; usually eight are admitted^: sable, M. zibellina; Japanese 
sable, 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 
the species of the subfamily. 

The greatest number of species are endemic to Eurasia (4). In 
America, there are two: the fisher, M. pennanti and the American 
marten, M. americana. 

Both in Eurasia, as well as in America, this genus has great 
practical significance, since in it are included very valuable fur- 
bearing species — all true martens and among them, the sable. The 
importance of the yellow-throated marten in this respect is not 
great. The kharza plays some role as a destroyer of game animals, 

^Reference to 10 species in the genus (Novikov, 1956), is 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: 1) 
sable M. zibellina 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 Identification of Species in the Genus Martens 

1 (2). Color patchy — there are portions of white, brownish-yel- 
low, blackish-brown and bright golden-yellow colors. Tail 
length constitutes about 2/3 of body length. Dimensions 
large — condylobasal length of skull of adults more than 

100 mm (body length of adults to 75-80 cm) 

yellow-throated marten, M. (Ch.) flavigula (p. 905). 

2(1). Coloration uniform — reddish-brown or brownish; a light 
spot may exist, on throat and chest only, white, yellowish 
or bright-yellow color, or head may be lighter than body, 
sometimes even whitish. Tail length comprises less than 
2/3 of body length. Dimensions small — condylobasal 
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 
than half the body length or equal to it — it hardly extends 
beyond the ends of the extended hind limbs. Light throat 
spot absent, or not clear, small and has irregular form and 
is not precisely outlined — not sharply demarcated from 
color of neighboring parts of body. Top of head usually 
lighter than back. Bony auditory bullae elongated and 
adjacent. Distance between them at middle of their lengths 
less or equal to half of distance from anterior point of 
bulla to posterior margin of lateral occipital process 
sable, M. (M.) zibellina (p. 757) 

4(3). Tail relatively long. Its length with terminal hairs more 
than half the body length — it extends beyond the ends of 



756 



507 



the extended hind limbs for more than 1/4 of its length. 
Throat spot variable in form, but well-defined and with 
sharp outline. Top of head same color as that of back. 
Auditory bullae shorter and widely separated. Dis- 
tance between them at middle of their length more than 
half of distance from anterior point of bulla and posterior 
margin of lateral occipital process. 
5 (6). Throat spot color pure white and usually gives off two 
posterior projections extending to the forelimb. Inner part 
of upper molar is not wider or only slightly wider than 
the outer (Figure 190). Tapering, 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 some- 
times digital pads^ stone marten, 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. Well-marked posteriorly pointing projection usu- 
ally found 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.). 

'Differentiation between stone and pine martens by form and color of throat spot, 
though most frequently practiced, is not always reliable (see section below "Descrip- 
tion" of species). For purposes of identification, characteristics of teeth structure are 
essential. 

* 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 

authors, more precisely: western Siberia north of Tobol'sk. 
1840. Mustela melampus. Wagner. Schreb. Saugeth. Suppl., 2, 

p. 229. Japan. 
1844. Mustela brachyura Temmink. Siebolds Fauna Japon. 

Mamm., p. 33. Japan, Hokkaido. 
1855^. Mustela zibellina var. asiatica. Brandt. Mem. mathem., 

phys. et natur., 7, p. 6, 23, Taf. 1, Fig. 1. Kamchatka.^ 
1855. M. zibellina var. alba. Brandt. Ibidem, p. 14, Taf. 2. Fig. 

5. 
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. Ibi- 
dem, p. 14. Taf. 3, Fig. 7. 
1855. M. zibellina var. maculata. Brandt. Ibidem, p. 14, Taf. 3, 

Fig. 9. 
1855. Mustela zibellina var. asiatica rupestris. Brandt. Ibidem, 

Taf. 2, Fig. 2. 
1855. Mustela zibellina var. asiatica sylvestris. Brandt. 

Ibidem, Taf. 2, Fig. 3. 
1918. "^Mustela zibellina subsp. kamtschadalica. Birula. Otchet 

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

^Date for individual volume — collection of works reprinted from volume VII of 
the "Memoirs" (see list of references at end of book). The pagination is given accord- 
ing 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, 
in Table 1, Fig. 1 is illustrated a normally colored sable, originating, as stated in the 
legend from Kamchatka. Formally, under such conditions, the name asiatica may be 
ascribed to Kamchatka sables. However, since in the whole sense of the paper it is 
clear that Brandt under his var. asiatica had all Asian sables this cannot be done. 

*In Russian original, not in chronological order — Sci. Ed. 



758 

1922. Mustela zibellina princeps Birula. Byalynitskii-Birula. 

Ezheg. Zoologich. muzeya. Ross. ak. nauk 1917-1921, 

22, p. 8. Barguzin range. 
508 1922, Mustela zibellina var. kamtschatica. Dybowski. Arch. 

Tow. Nauk. Lwowe, 1, p. 349, Nomen nudum. Kamchatka. 
1922. Mustela zibellina var. baicalensis. Dybowski. Ibidem, p. 

349. Nomen nudum. Vic. Baikal. 

1922. Mustela zibellina var. amurensis. Dybowski. Ibidem, p. 
349, Amur, Ussuri. 

1923. Mustela melampus coreensis*. Kuroda et Mori. Joum. Mam- 
mal. 4, p. 27. Tenan, South[em]. Chusei district]. Korea. 

1925. Martes zibellina yeniseensis. Ognev. Journ. Mamm. 6, 

No. 4, p. 277. Krasnoyarsk u[yesd], Yenisei guber[naya], 

taiga plain. 
1925. Martes zibellina sajanensis. Ognev. Ibidem, p. 278. 

Orzybai [=Orsyba] river, northern Sayan [mountains]. 
1925. Martes zibellina sahalinensis. Ognev. Ibidem, p. 279. 

Sakhalin, Vedernikovo. [=Saghalien, Wedernikovo]. 
1927. Martes zibellina correensis. Kishida. Choju Chosahokoku, 

4, p. 130. Korea. 
1927. Martes zibellina hamgyensis. Kishida. Dobuts Zasshi, 39, 

p. 509.** 
1941. Martes zibellina tungussensis. Kusnetzov. Tr. Mosk. 

zootekhnich. inst., I, p. 116. Lower Tunguskaya [river]. 
1941. M.iartes) z.(ibellina) sahalinensis arsenjevi. Kusnetzov. 

Ibidem, p. 122. Samarga River, Ussuri Territory. 
1941. M.{artes) z-iibellina) sahalinensis schantaricus . Kusnetzov. 

Ibidem, p. 122. Bolshoi [Great] Shantar Island, Okhotsk Sea. 
1943. ^Martes zibellina averini. Bashanov. Bazhanov. 

Kazakhskii fil. Ak. N. Souza SSR in 1942, 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 — Sci. Ed. 
**No type locality cited in Russian original — Sci. Ed. 
'Released apparently, in 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. Martes zibellina angarensis. Timofejev et Nadejev. 
Timofeev et Nadeev. Ibidem., p. 41. Angara valley in 
Boguchaev region, Krasnoyarskii territory. 

1955. Martes zibellina ilimpiensis. Timofejev et Nadejev. 
Timofeev et Nadeev. Ibidem., p. 44. Kochechumo water- 
shed — 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. 

1955. Martes zibellina obscura. Timofejev et Nadejev. Timofeev 
et Nadeev. Ibidem., p. 47. Upper Chikoi river — right tribu- 
tary of Selenga [river], Trans-Baikaliya. 

1956. "M.(artes) M.(artes) z.(ibellina) jakutensis. Kusnetzov 
(1941)". Novikov Khishchnye mlekopitayushchie fauny 
SSSR, p. 185. Yakutiya, Aldan Valley" (V.H.). 

509 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. 
Inner half of upper molar is wider than outer (V.H.). 

Description 

The sable, in its winter fur, is a very graceful and elegant animal. 
It 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 
and the claws are hidden in the dense hairs. The tail is compara- 
tively short, its length usually constituting about 1/3 of the body 
length and not exceeding half of its length. In the "skinned"* 
animal it is small (no more than 1/4 of its length) and extends 
slightly beyond the posteriorly extended hind legs. It is densely 
covered with long hairs and is very fluffy. The head of the sable 
appears very large, cuneate in form — the nasal part tapering, but 
slightly extended; the ears are large, upright, of triangular form 
with a broad base and a somewhat blunt tip. The eyes are bulging, 
quite large, black; the naked tip of the nose is black and the claws 
are black. 

Since the trunk of the sable is elongated, and the legs are 
relatively short, the animal always arches its back strongly, its 
hind parts are often higher than the shoulder; the tail is always 
fluffy. Nevertheless, the sable in its winter garb, is a very well- 
proportioned animal, an appearance which has nothing in common 
with a small cat. The general impression from the appearance of 
the sable confirms 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 the stones, 
fallen trees, etc. Finally, due to its broad, densely haired feet, it 
can confidently traverse loose snow without sinking in. The weight 
load of the sable track is less than in all our other species of the 
genus. 

The appearance of the sable in summer garb is entirely differ- 
ent — it looks thin and long, with longer legs and a thinner tail. The 
head seems disproportionately large, with huge ears, and the neck 
is thin and long. The relatively great breadth of the feet in the 
summer 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; 
the number of hairs in a 1 cm^ area of skin on the rump is equal 
to 600-700. 

♦Literally, "in flesh" — Sci. Ed. 



761 

The winter pelage of the sable is dense, with a large quantity 
of underfur, exceptionally fluffy and silky. The length of guard 
hairs on the croup equals 36-48 mm, the length of underfur, 24- 
28 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 20-25 underfur hairs; the total number of hairs in 
1 cm^ area of skin on the rump is, on the average, 13,500 
(Kuznetsov, 1951, Yenisei sables; Pavlova, 1951). In winter pelage 
the hairs of the tail 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, and facilitates the animal's movement on loose snow. 
510 While the winter fur of the sable is entirely variable in color 
(see below), the summer pelage of the animals in a single locality 
and in various parts of the range, is uniform. Even in very light 
races (for example, Tobolsk sable), the summer fur is very dark. 

Its color is 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 
they are somewhat darker than the color of the back. On the throat 
and along the lower neck, there is a lighter yellowish area with 
completely undefined outlines, spreading onto the sides of the neck. 
In some, it is absent. Between the ears and eyes, a dark area ex- 
tends to the nose; the cheeks, the region in front of the ears, the 
ears, and behind the ears is occupied by a light area of ochreous 
color merging posteriorly with the lateral parts of the throat spot. 
This pattern is sometimes absent or weakly defined, and the whole 
head is darker. The ear margins are trimmed with short ochreous 
hairs. 

The generally dark color of the summer sable is explained not 
so much by the coloration of the various hair categories as by the 
structure of the fur. The underfurs have a yellowish-brown color, 
the guard hairs and contour hairs are dark-brown or almost black. 
The previously mentioned ratio of 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 less intense color, but the contrast is insignificant. The 
tail has the same color as the back, but the terminal half is 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 con- 
siderably lighter than the remaining parts of the trunk. On the 
throat and neck there is 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 hav- 
ing either very monotone color, or more contrasting colors occur. 

The darker color of back and tail described above is character- 
istic of light-colored sables — the darker the sable, the less its con- 
trast, and in the darkest sables, not noticeable. So also with the 
color of the head — in the majority of cases, it is light-grayish and 
only in particularly dark sables is it the same color as the back; the 
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 
absent or only represented by a tiny, weakly colored part of the 
fur. Sometimes, on the contrary, the throat patch with all its 
variation in shape, is sharply outlined. The color of the patch is 



763 

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

A century of practice in the Russian fur business has elabo- 
rated a very precise categorization of color variation in sable fur. 
In it, no attention is assigned to the quality of the fur itself, and 
it is applied and adapted to all the geographical forms (sorts) of 
sable. These sorts are as follows: 

Golovka. Color particularly dark. All fur pitch-black or black- 
ish-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". Head dark. 

Skins of golovka are divided into vysokaya golovka (extra) — 
skin color pitch-black with almost pure black guard hairs; 
normalnaya golovka — skin color blackish-brown with brownish- 
black guard hairs. 

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) 
with chestnut hair tips. 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 — skin color dark chestnut with dark brown- 
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 tips. Head light, grayish. Throat patch large, 
bright. 

Skins of vorotovyi sable are divided into temnyi [dark] 
vorotovyi — skin color dark-brownish with weakly defined reddish 
tinges on sides, guard hairs chestnut, underfur with reddish-brown- 
ish hair tips; and normalnyi [normal] vorotovyi — skin color brown- 
ish with clear reddish tinges on 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 



764 

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

With regard to all remaining qualities, the skin of the mekhovoi 
sable is evaluated at about 15-20%, the dark vorotovyi — at 30- 
35%, and the vysokaya podgolovka — 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 correla- 
tion of color of different parts of the body, particularly the color 
tone of the head. 

Amplitude of individual variation in different populations and 
races may differ. In several parameters the range of variations is 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 con- 
ditions, well characterizes individual geographical races and 
populations (see below, section on "Geographic Variation"). In 
several races, separate types may rarely be found. Thus, in Tobol'sk 
sables, there are no golovka, and in some Trans-Baikal, light sa- 
bles 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 consider- 
able. 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 




Skin 


quality 






Total number 


Dark (%) 


Medium (%) 


Light (%) 


1941/42 


269 


16.4 


67.6 


16.0 


1943/44 


130 


19.3 


65.9 


14.8 


1944/45 


295 


8.8 


67.6 


23.6 


1948/49 


327 


29.0 


63.0 


8.0 


1949/50 


711 


29.5 


64.7 


5.8 


1950/51 


901 


11.3 


76.9 


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) — a ratio 
of 1:20. It 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.). How- 
ever, color is 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 still continues 
at the start 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 
great ("grizzled sable") or negligible, or completely absent 
("glukhaya sable"). White sables (albinos), chromistic and skew- 
bald (with white spots) may be found as exotypic variations. 
Sables with a bright orange tint as well as gray may also be en- 
countered. 

Sexual and age difference in the color of sables is absent. 
Young sables in their first autumn are clad in adult coats. 

The skull of sable is relatively narrow and elongated. Both the 
facial 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 por- 
tion. Zygomatic arches are relatively thin, the supraorbital proc- 
esses are well developed and broad, and the postorbital constriction 
is wide (more than the width of the skull above the canines). 

The mastoid processes do not protrude beyond the lower mar- 
gin 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 swol- 
len 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 cham- 
ber to the posterior edge of the paroccipital process. The auditory 
tubes are well developed. 
514 The longitudinal diameter of the upper carnassial tooth is 
approximately equal to the transverse diameter of the upper molar. 
In this latter the inner blade is considerably larger than the outer. 



766 





513 



Fig. 191. Skull of the sable, Maries (Martes) zibellina L. 



767 



Sexual differences in the skull are insignificant and represented 
only in the somewhat smaller dimensions of the female skull. Age 
variation is 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 supra- 
orbital processes), narrows. The interorbital width in males 8-9 
months old constitutes 18.5 mm on the average, while an animal 
older than 3 years is 20.2, corresponding postorbital constriction is 
18.0 and 15.5 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 skull by the time of sexual development of the 
animal. 

According to the character of these lines and other characters, 
the following age groups of sable are distinguished (Fig. 192; 
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 

Age I — young animals 8 to 10 months old, i.e. middle of first 
winter of life. The temporal lines separated 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 
slightly obliterated. Ratio of interorbital width to postorbital in 
males, 0.84-1.17 (M 1.06); in females, 0.78-1.14 (M 1.06). 

Age II — about 1 year, 8-10 months old (middle of second 
winter of life). Occipital crest well developed. There is a rudimen- 
tary sagittal crest, temporal lines come together posteriorly form- 
ing an acute angle. Incisors slightly worn. Ratio of interorbital 
width to postorbital in males, 1.04-1.22 (M 1.10); in females, 1.04- 
1.24 (M 1.11). 

Age III — about 2 years, 8-10 months old (middle of third win- 
ter of life). Occipital crest is moderately or strongly developed. 
Temporal lines come 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 (M 1.20); 
in 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 (M 
1.30); in females, 1.17-1.43 (M 1.24). 
515 The majority of the above-mentioned features, and above all, 

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 growth upward from both sides to meet each 
other. If in the young animal (Age II) this musculature is relatively 
small and muscles of the right and left sides 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 
forked 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 
ring. The length of the ossicle in Trans-Ural sables is 39.2-M 
41.4-43.2 mm (Yurgenson 1947)'^ Intestine length is 1000-M 

"Other features in the structure 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. 
Groups [A, B, C] are those given on in previous figure [Fig. 192] from Nadeev 

and Timofeev, 1955. 

1790-2700 cm, which constitutes 250.0-M448-630.0% of the 
length of the carcass without the hide (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, 320- 
510 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. 

Condylobasal length of skull of males, 74.1-94.6 mm; of fe- 
males, 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 mm; of females, 26.5-39.6 mm (after 
material of 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 it was determined 
according to the weight of the carcass without the hide. Its [hide] weight (n 230) is 
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 gm (Starkov, 
1947). The maximum known weight of a male is 1869 gm (V.H.). 

516 Systematic Position 

The sable is a typical representative of the genus Martes, closely 
related to the true martens; as with all of them, it is quite sharply 
distinguished from the fisher, M. pennanti. At the same time, the 
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 
and marten are accepted in what follows. 

It is possible that the American marten M. americana belongs 
to the species zibellina, although it 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 

This constitutes the overwhelming part of its range. It occupies a 
great part of the state territory — all the forested part of Siberia and 
the northern 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 dif- 
ficulties. In the European part of the USSR the sable was sub- 
jected 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 
it is met with under entirely different conditions — from 
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 
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 exam- 
ple, Minusinsk steppe). However, they themselves constituted ex- 
ceptions, 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 
expanses were formed in which the sable has completely disap- 
peared. Thus, the range was broken into separate parts. This was 
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 
persists. 

The separate areas inhabited by the sable became transformed 
into "islands", usually small, and in the majority of cases, espe- 
cially in the east, they are so completely isolated from each other 
that their natural restoration, given the cost of restocking, seemed 
517 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 was gradually taking place on a large 
scale, and in particular forest fires. 



772 

The picture of the range described at the beginning of the 20th 
century, was the result only of destructive human activity. As in- 
dicated by experiences in our day, the sable is quite viable and 
completely "contemporary" and, by no means, is going to become 
extinct, as some have attempted to portray it. 

The destruction of sable started so long ago, that, with the 
scarcity of historical data — above all, about Siberia — to restore the 
picture of its range is now quite difficult. Proceeding from the 
ecological characteristics of the sable, in regard to some territories 
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 
increased somewhat by the beginning of the second decade, during 
the period 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 sa- 
ble 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 
distributional data quite difficult. 

The history of the range of the sable is a clear example of a 
range change occurring under the influence of the anthropic fac- 
tor — at first, its deep, uncontrolled destruction, and then its planned 
restoration. Naturally, the range of sable cannot be fully restored 
to 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 outlines of the westernmost part of the range in the past 
(historical time) cannot now be reliably and completely restored. If 
there is more or less reliable information, not disputing the exist- 
ence of disagreements, along the Urals and the northeastern Euro- 
pean part of the Union, westward to Severnaya Dvina then the 
question of the occurrence of sable farther to the west is not so 
clear. Its discussion dates 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 de- 
nial. In recent years, however, new materials have been found 
(Kirikov, 1952, 1958, 1960) which reopened anew the discussion 
of this question'^ 
519 The localities farthest west for which there is 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 — the westernmost point; in the 16th and the 
first half of the 17th century). In Byelorussia, Minsk, Lutsk (now 
Volynsk district in the Ukraine; the southernmost point, about 51° 
N. 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, 
Polyakov, 1873, Sabaneev, 1875; Pleske, 1887; Ognev, 1931; Yurgenson, 1933, 
Zhitkov, 1937; 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. Data on the past and present distribution of the sable, Martes (Martes) 
zibellina L. in Europe. V.G. Heptner: 1 — concrete, and 2 — generalized mentions of 
the occurrence of sable during the 16th and 17th centuries; 3 — concrete, and 
4 — generalized information for the 18th century; 5 — concrete, and 6 — generalized 
information for the 19th century; 7 — probable northern and southern limits of the 
range in the northern European part of the country in the past; 8 — limits 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] at the end of the 17th 
century, where in 1834 it was still found somewhere in the north- 
ern parts of present Karelia (Kemsk county of Arkhangel'sk gov- 
ernance), and in the former Kholmogorsk county, and in the 16th 
century, in Velikii Ustyug district. This is the southernmost place 
of occurrence of sable in the section of the northern European part 
of the USSR under consideration. 

Concerning Finmark, the sable is mentioned in Scandinavian 
folklore in the 9th-10th centuries. Finally, northern Finland and 
northern 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 
the west occupied the taiga of Arkhangel'sk district, Kareliya, and 
the 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 sable was, apparently, widely distributed along the Mezen' and 
its western tributary, the Vashka (Udor), i.e. evidently also in the 
interfluve between it 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 
of its range reached to the mouth of the Ob' river. 

In the Vychegda basin, sable existed and were still quite com- 
mon at the beginning of the 17th century, being found not only 

(Fig. 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 — only 
individual peripheral points. 

'Чп the manuscript of the monk Epifanii — a minor companion of Protopop 
Avvakum whose personal report of the place is ascribed to one hunter, who said: 
"would it be any wonder, if ... 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, Vym' and others), but also along its left 
(southern) tributaries — the Sysola as well as Bolshaya and Malaya 
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 even farther to the west. 

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 Kama to the northeast of 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 
520 of the former Vyatsk governance (Sabaneev, 1875) is supported by 
documented places of occurrence; however, concerning more west- 
ern 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) and along the southwestern slope of the 
Urals to the east of the city of Ufa even in the first half of the 19th 
century (Sabaneev, 1875)'^ 

The true southern border of sable distribution in the Urals in 
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 is the result of secondhand rewritings and simplification of 
original sources. Pallas (1786) writes: "It 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. 



777 

region of 55-56° N. lat. — the sources and upper reaches of the Ufa 
(Pallas, 1786) and Kaslinsk mountain (Sabaneev, 1875)^^. 

In summarizing everything said, it is possible to accept that, in 
historical time, and, still in part at the end of the 18th and even the 
19th centuries, the southern boundary of the range of sable in the 
European part of the USSR began in the Urals at about 55-56° N. 
lat., crossed the Ufa in its upper or middle course, extended to the 
Kama somewhat above the mouth of the Belaya, passed thither to 
the upper Vyatka and farther to the region of Velikii Ustyug. 
Thence, it turned directly westwards to Olonetsk territory (south- 
ern Kareliya), passing somewhere through the region of Lake 
Onezhsk [Onega], possibly through its middle, or even southern, part^. 

Beyond this line, it is possible that separate areas of 
occurrence were found in Lithuania and Byelorussia. It is worth 
noting, 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 inhabi- 
tation, isolated (artificially?) from the main range was found in the 
southwest, similar to several of the present Siberian outliers. The re- 
ality is that it lies to the south of the southern border of Neolithic sable 
fossil finds as they are known to us at the present time (see below). 

The northern border of the range extended, apparently, along 
the northern border of the forest — in the west, Kola was mentioned 
and in the east — Pustozersk, and along the border of the forest 
lying to its east, i.e. 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 5Г 
N. lat.", is 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 
postulate the existence of sable south of 55-56°, is now only possible by analogy to some 
other northern species which descend along the Urals south as far as 52°, and for very 
remote times. Evidence concerning 5Г, accepted by many authors after Sabaneev, is based 
on misunderstanding, — 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 

in Europe agrees well with the limits of the range in the Neolithic, 
which generally corresponds to the outline of the [present] south- 
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, pro- 
ceeds somewhat more to the north of Kasimov, crosses the Oka 
[river] above Murom, the Sura in its lower reaches and the Volga 
slightly above the mouth of the Kama. This recent boundary crosses 
the Vatka at about 50° E. long. — in its lower reaches and again 
crossing the Kama above the mouth of the Belaya, reaches the 
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 rutilus, C. 
rufocanus), wood lemming (Myopus schisticolor) and in part 
Siberian chipmunk (Tamias sibiricus) extend very far to the west 
in the north of our country — all, except the chipmunk, to Scandi- 
navia. 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- 
tributed far to the west in the past^^. 

The present (50' s) western limit of the range of sable takes the 
following form. In the north, it begins slightly to the north of the 

"Middendorf (1867) affirmed that sable never existed to the west of Syktyvkar 
(Ust'-Sysol'sk). 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 
penetration of sable to the west, but doubted it and only Pleske (1886) believed in the 
existence of sable on the Kola Peninsula still in the 18th century. 



779 

sources of the Lyapin at approximately the latitude 63° 30' N. lat. 
(a little south of the limit of nut-pine) 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 re- 
gion (V.Ya. Parovshchikov). 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 
Obsk Bay. In any case, this line included the basin of the right 
tributary of the lower Ob', the Kunovat river (65° N. lat.). Along 
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 (left tributaries of the Yenisei); crosses the Yenisei some- 
522 where in the Dudinka region and rises along the right bank of the 
Yenisei to the region of Lake Kit (of the Noril'sk group) and upper 
Noril'sk creek. It falls only slightly short of Noril'sk and here at- 
tains a latitude of 68° 40' N. (Podarevskii, 1935) or even includes 
Lake Pyasino and the upper Pyasina at 70° 30' N. (Kirikov, 1960). 

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 Olenek— at 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, probably about 68", includes the basins of the 
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 en- 
compasses not only the Omoloi basin, but also the Bol'shoi and 
Malyi Anyui (Middendorf, 1867). From the Anyui, the range bor- 
der 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). 

The description given of the northern border of sable range in 
Siberia was constructed on the basis of direct evidence of its con- 
temporary distribution and its distribution in relatively recent past. 
It is, however, well founded (archival material dated from the 16th 
century; Kirikov, 1958, 1960), that in the expanse from the Ob' to 
the Kolyma, the actual natural range boundary ("reconstructed") 
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 to the reconstructed [border]. In some 
places on the northern border, the sable also lives outside the [taiga] 
forest zone — in krumholz, in deciduous forests of river valleys 
(Anadyr') and in nut-pine thickets (Koryatsk land). 

The shore of the Pacific Ocean constitutes the eastern range 
boundary. Within the range are Shantar and Sakhalin islands. Of 
the Kuril islands, the sable is shown to have resided on Iturup and 
Kunashir (Snou, 1902; Voronov, 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 
absent, as on the Commanders. 

The southern border of the reconstructed sable range began in 
the Urals in the upper Ufa at a latitude 55°-56° (see above) and 
entered Siberia near Chelyabinsk (Pallas, 1786). Further to the 
east, the border passed through the whole of eastern Siberia in the 
forest-steppe zone. From Chelyabinsk, it 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 



781 

reached the Barabinsk forest- steppe. Here the range boundary went 
to the mouth of the Om, apparently, north of Lake Ghana and 
reached Lake Ubinskoe and to the Ob' at Novosibirsk. This is the 
border in the 17th century, and, in some places, the 18th century 
(Pallas, 1786; Sabaneev, 1875; Ognev, 1931; Kirikov, 1960). 

From Novosibirsk, the boundary turned sharply to the south 
and passed towards the foothills of Altai, apparently in the Biisk 
region. Here, the border turned back to the west and then to the 
south and southeast, embracing the Altai. In this region, the bound- 
ary again passed along the edge of the foothill forests at the lati- 
tude of Biisk; farther, it passed around the Tigeretsk range to the 
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 bound- 
524 ary included the Kurchum and the region of Lake МагкакоГ 
(Ognov, 1931; Yanushevich and Blagoveshchenskii, 1952; 
Afanas'ev et. al., 1953; V.G. Heptner). At МагкакоГ, the bound- 
ary of the range reached the state border". It is probable that the 
Altai portion of the sable range was somewhat broader. It may be 
that sable lived also in the ribbons of forest in the Cis-Altai steppes 
and extended south to the Irtysh. However, there is no information 
on this point. 

In the expanse from the southern Altai to the Pacific Ocean, 
the sable range reaches the state border (Tannu-Ola, for instance) 
or beyond it. In Ussuri Territory, the sable is known as far south 
as the Kedrovaya river (Kedrovaya Pad' preserve) between 
Vladivostok and Pos'et (Ognev, 1931), i.e. 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° 
N. lat. 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 
the left bank of the upper Irtysh and across the Alakol'sk lakes (!) it extends to Issyk- 
Kul'. 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 
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 Sibe- 
rian 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 compared to the beginning of the current century, it still 

526 remains fragmented into separate parts, some massive and some 

relatively small. The regions inhabited by sable are as follows 

(Fig. 196). 




Boundrles of 
^ "Centers" of 

ccurrence of sable 



524 Fig. 196. General scheme of recent range of the sable in the USSR. Figures designate 
the separate "centers"of occurrence. Data up to 1950 (after Nadeev and Timofeev, 

1955, simplified). 

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




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785 




526 Fig. 198. Boundary of sable distribution in the extreme west of the range — in the 
Ural-Ob' region of occurrence (after Nadeev and Timofeev, 1955). 

Ussuri, southern Trans-Baikaliya and Altai. Between these centers 
are interspersed, still smaller and sometimes very small regions of 
sable occupation, sometimes lying very far from each other, as in 
Yakutiya (after Nadeev and Timofeev, 1955; the names of the 
separate centers do not correspond to those given by these au- 
thors). 



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 Ulan- 
Bator), 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 finger of the range which extends south along the Mongo- 
lian Altai, apparently transgresses somewhat into Chinese 
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 Barga steppe and the plains of the former 
Central Manchuria. It exists in the Bol'shoi 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 in the 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; 
Lukashkin and Zhernakov, 1934 and others). Details of sable dis- 
tribution 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 in sable is well marked, and if taken in 
the sense of morphologically extreme forms, the amplitude of vari- 
ation is quite significant. This variation is expressed in general 
body and skull dimensions (its proportions are nearly invariable) 
and, particularly, colors. Fur quality varies considerably: its 
density, fluffiness, softness and fineness. 

Geographic variations in sable have been noticed and employed 
practically in the fur trade for a long time, where they were 




^^ Distribution in 1935 (Цр Distribution in 1950 ^ Places of sable introductions 

527 Fig. 200. 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 and in its 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 
(Ognev, 1925, 1931), although individual reports appeared earlier 
(Birulya, 1916, 1918) and attempts at analysis were made more 
than 100 years ago (Brandt, 1855). The latest 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). 

^••At the International fur auction in Leningrad in 1961, the highest lot price of 
sables of various grades was as follows: Tobol'sk — 28 dollars, Altai — 30 dollars, 
Amur — 31 dollars; Yenisei — 41 dollars, Tuvinsk — 67 dollars, Kamchatka — 190 dol- 
lars, Yakutsk — 300 dollars, and the Barguzinsk — 405 dollars per skin hide (B.A. 
Kuznetsov). Ratio, 1 : 14.4. This example is a utilitarian demonstration of the reality 
of geographic race. 



789 




528 Fig. 202. Reconstructed species range* of sable Martes (Martes) zibellina L. Dashed 
line designates region in Lithuania and Byelorussia relative to which literature data 
exist on occurrence of sable in the past. V.G. Heptner. 

In the sable very wide individual variation in color is charac- 
teristic (see above), and therefore the study of its geographic 
variation is a very difficult task, and racial characteristics are some- 
times ill-defined. At the same time, analysis of a mass of collected 
material, 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 
percentage of various color types. 

*Japanese populations south of Hokkaido are now considered a separate species, 
M. melampus — Sci. Ed. 



790 

In the literature, it was frequently suggested, mainly on the 
basis of information from hunters and traders, that sable color in 
one narrow region changes depending on the land inhabited (nut 
pine — broad-leaved forest), altitude (valleys — mountains) and other 
ecological conditions, and even annual character of food. Zoologi- 
cally these observations have not been confirmed; however, to 
completely deny a connection between characteristics and known 
ecological conditions is not suitable. This is clear, for example, 
from those 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 
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 zoology, especially in a stricter and broader interpretation of 
this concept. 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 will, undoubtedly, be combined. 

On the other hand, it is necessary to have in mind some special 
conditions of existence in the sable. This animal is an animal dis- 
tinguished by its great attachment to place within a quite strictly 
delineated locality. Its range under natural conditions, was always 
broken into separate more or less isolated parts, especially in the 
east. In the past century, in connection with its extermination, it 
was divided into small parts, surviving — in places for a long time — 
without connection with each other. All this may have influenced 
the formation of small local populations. Easiest of all, in this 
530 sense, change in the ratio of color types can be expected. In some 
cases, separate names were noted, apparently, for local population 
("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 
others (see below). The reality of 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 
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, it becomes coarse. This 
manifestation of general regularity is mentioned by furriers — the 
coarsening of the fur in more humid regions, especially in districts 
with a maritime climate. The darkest sables inhabit the central part 
of the range and the lightest — its margins (Ob' lowland, Sakhalin, 
Shantar islands). In montane regions, sable are darker than on the 
plains (Kuznetsov, 1941). 

As is clear 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 
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. 

Body length of males, 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- 
sive original data and literature). 

Condylobasal length of skull of males (159 samples) 79.7-M 
85.7-90.6 mm, females (87) 73.8-M 78.9-83.8 mm; zygomatic 



792 

width of males, 44.1-M 48.0-54.8 mm, females 39.0-M 43.6-47.2 
mm; height of male skull 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 consti- 
tutes 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.) z- averini Bashanov, 1943 (synonym 
altaica). 

531 Body and skull dimensions comparatively large, color quite 

dark and varies from sandy-yellow to dark brown. 

Golovka and Podgolovka sorts 26%, Vorotovaya 66% and 
Mekhovaya 8% (n=1000). 

Found in Altai, particularly Bukhtarma basin, northeastern part 
of Tuvinsk ASSR (Kemchug basin). 

Outside USSR, found in southern slopes of Mongolian Altai 
(Bulugun river). 

In several places in Altai, Barguzin sable was introduced, yield- 
ing hybrids with local sable. 

4. Yenisei sable, M. (M.) z. jeniseensis Ognev, 1925. 
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 light, but noticeably darker than 
Tobol'sk and Yenisei. 

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 in Sayans. 

Outside USSR, found in Kosogol lake district in Mongolia. 

7. 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, 
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%, 
and Mekhovaya 31% (n = 2600). 

Found north of Nizhnaya Tungusska to northern border of 
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) . 

Dimensions relatively small — one of smallest races of species; 
color entirely dark, but overall somewhat lighter than Vitim sable. 



794 

Body length of males, 394-M 415-420 mm, females, 360-M 
382-410 mm; tail length of males, 122-M 144-155 mm, females, 
120-M 129-145 mm; condylobasal length of males, 73.9-M 77.9- 
81.0 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 29.7- 
31.4 mm. 
532 Golovka and Podgolovka sorts constitute 45%, Vorotovaya 46% 

and Mekhovaya 9% (n=786). 

Found on eastern shore of [Lake] Baikal; Barguzin range from 
right tributaries of 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 
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] in Mongolian Republic. 

Extremely valuable form. 

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 42% 
and Mekhovaya 5% (Kuznetsov, 1941). 



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 sorts constitute 4%, Vorotovaya 59% 
and Mekhovaya 47% (Kuznetsov, 1941). 

Found in Sakhalin [Island]. 

Absent outside USSR. 

It is probable, the name brachyura belongs to the form since, 
according to some data, 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 sorts constitute 16%, Vorotovaya 67% 
and Mekhovaya 17%. 

Found in Ussuri Territory, mainly in Sikhote-Alin' taiga. 

Outside USSR, found in contiguous parts of North-east China. 

Relationship of this form to form coreenis {hamgyensis) found 
in Korean Peninsula needs clarification. 

15. Shan tar sable, M. (Л/.) z. schantaricus Kusnetzov, 1941. 
Dimensions as in Ussuri sable. Color darker than Sakhalin and 

Ussuri. Pelage quite fluffy. 
533 Golovka and Podgolovka sorts constitute 32%, Vorotovaya 63% 

and Mekhovaya 5%. 

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, M 152-172 mm; condylobasal length of male skull, 
85.6, M 90.4-94.6 mm (M 88.5 mm-93), females, 79-4, M 82.1- 



796 

84.4 mm (М 80.8 mm-76); zygomatic width of males, 51.3, M 
53.5-56.2 mm (M 50.2 mm-81), females, 43.6, M 46.3-52.8 mm 
(M 44.9 mm-74); skull height of males, 31.6, M 33.5-35.1 mm (M 
33.3 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). 

Golovka and Podgolovka sorts constitute 16.0%, Vorotovaya 
76.0% and Mekhovaya 8.0%. 

Found in Kamchatka and Koryak land in the Apuka basin. 

Absent outside USSR. 

17. Kuril sable, M. (M.) z. brachyura Temminck, 1840. 

Diagnostic features of sable found in the Kuril Islands are not 
sufficiently clear. Some authors relate its name to the named Japa- 
nese form. Apparently, it belongs to the Sakhalin in sable type. 

On the whole, all forms enumerated may be regarded as aggre- 
gating into seven groups (large populations): Tobol'sk — Pechora, 
Urals and West Siberia to Yenisei; Altai [and]; Sayan — Tuvinsk 
ASSR, Sayans and north to Kuznetsk Altai; Yenisei — right bank 
of Yenisei to Lena; Trans-Baikaliya — south to border of species 
range, east to watershed of Zeya and Bureya, north, including Vitim; 
Sakhalin — lower Amur, Sakhalin, Ussuri Territory and Shantar Is- 
lands; Kamchatka — Kamchatka and Koryak land (B.A. Kuznetsov). 

Weight of sables from various parts of the range is as follows 
(Kuznetsov, 1941): Yenisei sable— males (93) 1030, M 1307-1500 
g, females (125) 880, M 1182-1250 g; Sayan sable— July males 
(22) 1200, M 1433-1800 g, females (21) 870, M 980-1300 g, 
January males (22) 1110, M 1472-1810 g, females (21) 820, M 
1132-1560 g; Sakhalin sable— July males (22) 940, M 1239-1600 
g, females (25) 800, M905-1130 g, January males (22) 880, M 
1193-1500g, females (25) 740, M 916-1120 g; Kamchatka sa- 
ble—males (4) 800, M about 1500-1869 g, females (5) 600, M 
about 1100-1140 g. 

Outside the limits of the USSR, the following forms are noted 
(sometimes, several of them are related to "Л/. melampus"): 1) M. 
z. brachyura Temminck, 1844 — in Hokkaido, Japan; 2) M. (M.) 
z. melampus Wagner, 1840 — in Honshu (Hondo), Shikoku, 
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.). 



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 con- 
tinuously increasing, as a result of the system of state measures for 
534 protection and restoration of this very valuable fur-bearing animal 
and they far exceed the prerevolutionary. 

At the present time, there are many regions in Siberia where 
the population density of sable attained a level which previously 
seemed impossible and was not actually present. The latter is ex- 
plained in that nowhere in the past within the range of the sable, 
were there territories 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 al- 
lowed 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 numbers were 
begun at different times. Therefore, at present the number and 
density of sable still do not correspond everywhere to the capacity 
of taiga areas and its distribution is not the result of natural regu- 
lation. Evaluated on the results of the hunting harvest, the total 
number of sable in the 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, 
Kemerov, Irkutsk, 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 
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 ex- 
ploited — 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 1.5- 
2.0 sables per 10 km^ with 8.3-8.5 track encounters per 10 km in 
good areas (forests of white birch [B. alba], flooded forests) — 1.4 
(6.8-7.3 track encounters), in moderate (broad-leaved and fir for- 
ests) — 1.3 (4.8 track encounters) and in the worse — 0.1-0.3 (0.8- 
1.4 track encounters). The total number of Kamchatka sables is 
estimated at 19,000 (average catch is 7,650 individuals, equaling 
an average growth of 40.3%; A. A. Vershinin). 

In northwestern Yakutia, one sable has 15-26 km^ and taking 
into consideration uninhabited regions — 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 
forests of the taiga type. 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 
areas, and cross their boundaries only during times of migration. 
Preference is shown for dark coniferous plantations almost always 
having in their composition Siberian stone pine. This is because of 
the great availability of food (pine-nuts, forest voles) and the pro- 
tective nature of this plant formation. 

Only as a result of irrational utilization, sable became mainly 
inhabitants of montane taiga, rocky talus and frequently even 
montane balds. At the present time, in the same way as in the 
distant past, sable willingly occupy and attain considerable density 
where lowland pine forests and swamps predominate, although even 
here, they prefer the islands of nut-pine and spruce forests, mar- 
gins of moss bogs and narrow belts of dark coniferous forests 
535 along river banks. Sable also lives under unique conditions such as 
montane forests of Erman's birch in Kamchatka, in larch forest 
with an understory of green alder [Л. viridis] in the Zhigansk 
region and even in forest tundra in the Gizhiga and Penzhina re- 
gions. 

The close linkage of sable with the Siberian nut-pine is exclu- 
sively conditioned by the general scarcity of food available for it 

*Local word — Sci. Ed. 



799 

in the Siberian taiga. This connection is not an indispensable con- 
dition of its existence. 

Food. The sable is a polyphagus carnivore. In addition to warm- 
blooded animals, plant food — seeds of Siberian and Korean nut- 
pine and prostrate nut-pine, berries and insects are of great 
significance in its food. The composition of its food varies greatly 
in various places. For example, differences in food type of lowland 
and montane sables are quantifiable. As in pine marten, sable foods 
reveal geographical, stand-type and sexual differences. 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, 
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, Khabarovsk Territory and a 
series of other regions, only two elements — forest voles and pine- 
nuts — or the forest voles 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 ptarmi- 
gan, root vole, "nuts" of prostrate nut-pine and fish are character- 
536 istic foods. Red-backed voles, pine "nuts" and mountain ash 
predominate in foods of the Barguzin sable. For Kondo-Sos'vinsk 
sable, capercaillie and, in general, tetraonid birds are typically the 
most often encountered, and for Pri-Amur sable — forest voles and 
cowberry. 






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 de- 
pending 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 utilization of second- 
ary foods, or those which are not regularly utilized. 
537 Food composition is not the same even between small 
populations living in separate montane valleys in one region. This 
is 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 


Bol'shaya river 


Davshe river 


Tarkulik river 


Mouselike rodents 


90.0 


75.0 


92.3 


Birds 


13.0 


7.5 


3.8 


Reptiles 


12.2 


5.0 


3.8 


Insects 


23.6 


25.0 


15.3 


Pine "nuts" 


30.0 


55.0 


73.0 


Berries 


16.9 


20.0 


3.8 



Food composition of sable changes in snowy and nonsnowy 
periods of the year in various parts of the range (Tables 50, 51). 

The seasonal 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], wood lemmings [L. 
schisticolor], shrews 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 — when they spend the night under snow. 
In winter periods with deep snow, sable feed mainly on berries of 
mountain ash, and in autumn, or when there is little snow — on 
cowberry, and in summer, on blueberry. Deep snow makes it dif- 
ficult to obtain voles, pine "nuts" and some other food. 



802 



Table 50. Geographic variation in sable foods in the snow-free period of the year 

(occurrence in%). 



\ Region, 


West 


Northeastern 


Barguzin 


Middle 


\ Number of 


Siberian 


Altai 


preserve 


Sakhalin 


\ items, 


lowlands 








\ Investigator 










\ 




76 


314 


357 


Food \ 


Raevskii, 


Shaposhnikov, 


Timofeev 


E. Volkova 


composition \ 


1947 


1956 


1948 




Forest voles 


57.7 


36.8 


73.5 


64.0 


Siberian chipmunk 


— 


3.9 


3.5 


5.0 


Eurasian flying-squirrel 


— 


1.3 


2.5 


1.0 


Common squirrel 


7.7 


3.9 


— 


0.2 


*Pika 


— 


0.5 


1.9 


— 


Shrews 


7.7 


19.7 


13.0 


2.0 


Birds 


23.1 


1.0 


6.9 


25.0 


Tetraonids 


7.7 


3.9 


1.2 


No information 


Hazel hen 


7.7 


— 


1.2 


No information 


Bird eggs 


3.8 


1.3 


0.6 


0.6 


Insects 


53.8 


13.1 


5.7 


15.0 


Pine "nuts" 


3.8 


36.8 


37.6 


34.0 


Berries 


44.5 


13.1 


35.7 


66.0 



*The Russian word senostavka ("haystacker") is used, rather than the more com- 
mon pishchuka — Sci. Ed. 

Sable eat carrion and the food remains of other carnivores — 
this explains, for example the occurrence of squirrel in the food of 
sable (Shaposhnikov, 1956; P.P. Tarasov). 

Utilization of the Eurasian flying-squirrel [Pteromys volans] is 
associated with diurnal activity of sable; the latter can overtake it 
in its holes only during the day (the flying squirrel is a nocturnal 
animal). 

In Kamchatka, the frequency of the most important food com- 
ponent — voles — varied in different years from 35% to 84% (A. A. 
Vershinin) (Table 52). 

Summing up the factual data on sable foods, one may affirm 
that the significance of the Siberian chipmunk in its food is ex- 
tremely limited and is not found everywhere: in summer, its occur- 
rence in sable food does not exceed 3-5% and in winter, is even 
less. 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 



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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 


69.8 


66.6 


Voles 


61.1 


61.0 


Root voles [Microtus oeconomus] 


24.0 


24.0 


Red-backed voles {Clethrionomys 


rutilus] 20.4 


12.7 


Red-gray vole [C. rufocanus] 


16.5 


19.4 


Squirrel [Sciurus vulgaris] 


0.8 


0.4 


White hare [Lepus timidus] 


2.7 


2.4 


Pika [Ochotona hyperborea] 


0.4 


— 


Birds 


20.8 


6.1 


Capercaillie [Tetrao parvirostris] 


1.7 


1.2 


Willow and rock ptarmigan 


1.5 


1.2 


Small birds 


12.5 


3.6 


Bird eggs 


1.1 


— 


Fish 


18.1 


11.9 


Salmon 


10.2 


— 


Insects 


2.5 


— 


Hymenopterous insects 


0.9 


— 


Plant food 


53.5 


15.2 


"Nutlets" of prostrate nut pine 


10.3 


2.7 


Berries 


— 


12.5 


Mountain ash [Sorbus] 


21.6 


3.6 


Hawthorn [Crataegus] 


20.9 


6.1 


Chokecherry [Prunus sp.] 


4.8 


— 


Rose [Rosa sp.] 


7.3 


— 



Studied. The percentage of squirrel occurring in winter foods of 
sable ranges from 0.2 to 11.3%, averaging 3.5%. In the summer 
period, it ranges from 0.2 to 7.7% (on average 2.9%). 

As a result of data communicated on the significance of squir- 
rel in sable nutrition (Timofeev, 1962) it was revealed that in various 
parts of the range and in various years, the percentage of occur- 
rence of squirrel in stomachs and extrement of sable varied from 
and 26%; on average, 4.4%. Of 33 data series, the percentage of 
squirrel occurrence 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 number of 
squirrels is 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 sable cannot penetrate into the burrows of pika and chip- 
munk due to its large size. In winter, the chipmunk sleeps deep 
inside its burrow, and even the brown bear cannot dig it out. 
However, in summer sables catch chipmunks by ambush in the 
same way as it does with pika. In winter, pikas come out of their 
burrows with their hay stores under the snow where they are some- 
times overtaken by sable. Sable catch tetraonid birds in their snow 
burrows, but this does not occur often. In the north Urals, the sable 
540 and kida catch one tetraonid for every 10-15 attempts, when the 
birds succeed in flying away. 

In captivity, when offered 200 to 300 g food, sable did not eat 
it all, but hid some (Dul'keit, 1929); a norm of 170-180 g and 
150-160 g of food were sufficient for males and females respec- 
tively. 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 be- 
hind, 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 Kamchatka and in different years (A. A. 
Vershinin). 

The sable does not make food caches. In the presence of large 
prey, it makes a temporary lair nearby. 



806 

Home range. In August, the sable litter disperses. Young 
sables begin to wander, searching for a permanent home range. 
Food shortage intensifies their wandering. When the period of 
settlement is completed, sable become strictly sedentary animals. 
The home range selected is usually used for many years, often 
until death. Only forest fire, felling of the forest and similar events 
make the sable leave its home range. After the death of the "owner", 
another one occupies the range, preserving its boundaries which 
are determined 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 
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 as though the reflex of range 
defense is lost. Competition is not intensified, but migration 
appears, mainly among juveniles — the less settled part of the popu- 
lation (O.K. Gusev"). 

The size of the home range is determined by the abundance of 
food, the presence of escape and nesting conditions, and also the 
percentage correlation inside the home range area of various quali- 
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. Some- 
times, 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 
americana] length of daily movement in the USA is equal to 8.4 
541 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 Sayans south of 
Krasnoyarsk). Place of day-time lair of sable in areas where Barguzin sable was 
introduced. Photograph by G.D. Dul'keit. 

these ranges, one animal may occupy a home range of 2590-3885 
ha (states of Idaho and Montana). The size of the daily range of 
American marten in Glacier National Park (Montana) is 233 ha on 
average for males (77.7-440.3 ha) and for females — 69.9 ha (7.7- 
181.3 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 
(Yakutiya), home range comprises about 30 km^ (Romanov, 1938). 
In northwestern Yakutiya (Tavrovskii, 1958), 5 out of 8 sable trails 

*Correct citation is Hawley, V.[ernon], 1957 — Sci. Ed. 



808 

did not exceed a circle 4 km in diameter (about 12.6 km^), in 
eastern Sayan — from 4 to 10 km^ (Kozhanchikovy, 1924), on the 
northeastern shore of Baikal — about 10 km^ (Doppel'mair, 1926), 
in the upper Katun' river (Altai) — from 8 to 12-15 km^ (V.V. 
Dmitriev and P. Bashilov), in the former Kondo-Sosyvinsk pre- 
serve, on average about 10 km^ (Raevskii, 1947), and in the north 
Urals on the upper Ilych river — about 4-6 km^. In the Eastern 
Sayan foothills, in case of a moderate crop of principal food, a 
home range of 2.5-3.0 km^ is sufficient for one sable (Dul'keit, 
1957); the maximum density in this case was 1 sable per 2.6-2.7 
kml 

Within the limits of the distribution that are observed, conse- 
quently, there is fluctuation in size of range from 4 to 30 km^. 
Such fluctuations occur locally and in limited territories. 
542 Burrows and shelters. Breeding, winter, autumn and tempo- 
rary 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 
lying on the ground, and quite rarely — in holes in the ground and 
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 
(Shaposhnikov, 1956) and in Primor'sk Territory (Bromlei, 1956), 
they are not manifested. Permanent winter nests show signs of 
significant occupation (ice-covered entrances, excrement, 
ectoparasites and others). 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 
are of various forms. Within the boundary of the home range, the 
sable has some of these. In the Eastern 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). 

Daily activity and behavior. The sable belongs among those 
animals with an indefinitely defined rhythm of daily activity. Indi- 
vidual sables are observed with crepuscular-nocturnal — others with 
diurnal types of activity. It is not even rare that one and the same 
animal is active during the day, and then in the twilight and at 
night, depending upon the success of its hunting, weather condi- 
tions 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 
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 — crepuscular- 
noctural. On Blol'shoi] Shantar Island and in the northeastern Altai, 
the animals have no definite rhythm. In the foothills of the Eastern 
Sayan, in 68% of the cases, sables hunt during the day (Dul'keit, 
1957). 

The sable is a typical terrestrial carnivore. On trees, it does not 
feel self-confident, it climbs more poorly than pine marten and is 
not capable of moving 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 



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 
about freely there and capturing food items. 

The senses of hearing and smell are well developed. Vision is 
weak and monochromic. The sable swims well. 

In its 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 avail- 
ability 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 tires. The small size of the female leads to the fact 
that with the same number of jumps and energy utilized, the fe- 
male runs a shorter distance than the male. 

In the basins of Konda and Sos'va rivers (Raevskii, 1947), the 
sable moves 2-3 km per day (early in spring, 4-5 km); in the 
northern Urals (Ivdel'skii region) — 8-9 km; m northeastern Altai 
(Shaposhnikov, 1956) — from 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 km (Afanas'ev, 1961). For the 
Eastern Sayan (Solov'ev 1921), the path is indicated to be about 
5 km; in Barguzin preserve — about 10 km on the average; in 
Kamchatka (Kazarinov, 1946) — about 12-15 km; in the Sikhote- 
Alin' range (Bromlei, 1956) and in snow 12-14 cm deep — from 



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 1953- 
1955 period did not exceed 9.2 km. Average length of the daily 
trail was 5.6 km for males, and for females — 4.5 km. Average 
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 km per 
day, and in the second half of winter — 5.9 km. The larger the 
sable, the longer its trail per day. Deep snow and frost shorten it, 
544 as well as abundant food; in years of nut pine harvest, the sable 
sometimes becomes satiated in a 1-km journey. In northwestern 
Yakutiya (Tavrovskii, 1958), the average length of the daily jour- 
ney 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 character- 
istic directional integrity. In winter, the sable catches mouse-like 
rodents under the snow; on warmer days, they often ambush them 
on the surface of the snow like a cat (northeastern Altai: Timofeev, 
1951; Shaposhnikov, 1956). It catches tetraonid birds in the snow 
holes where they spend the night. It searches for nut pine "nutlets" 
and cowberries under the snow. Mountain ash berries are eaten 
directly from the branches. 

Seasonal migration and transgressions. The sable is a seden- 
tary animal, but translocations do occur in the species. Above all, 
there is the yearly dispersal of young in August to a new place, 
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 de- 
gree, a nomadic or seminomadic mode of life. They roam very widely 
within the limits of their range in search for the most available foods. 

For Kamchatka (Vershinin and Dolgorukov, 1947) seasonal 
migrations are noted above all; annually, in the second half of 
winter, sables descend 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. However, north Ural sable hunters believe that the sable 
dies in its place and does not leave the region of its inhabitation. 
Apparently, sable behavior is 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 few sables are 
saved by fleeing; the majority of them die from the fire. Appar- 
ently, massive migrations due to food shortage is possible. In the 
Sayans, they were noted in 1875, 1888, 1900, 1915, etc., approxi- 
mately every 12 to 15 years. This question is not yet well studied. 

Sables tagged in the northern Urals were caught at distances of 
2.0; 2.0; 4.0 and 4.5 km 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 
length of daily trails of dispersing sables reaches 10 km and more, the 
animals moving with longer jumps than during the fattening period. 
With increase in snow depth, dispersal stops. Areas from which sables 
disperse are usually those which are not fully exploited. They settle 
in areas where sables were removed during the harvest period. 

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 
not less than two times, and one male 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 — 
545 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. 
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 

The least percentage of infertility 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- 
tological investigations, become pregnant in the second year of life 
(Zaleker, 1953). The percentage of pregnant females among 
Barguzin sables in the second year of life in nature was from 26 
to 33% (Baevskii, 1956). In adult females, it ranged from 50 to 
72.9% depending on nutritional conditions in the data year. In 
regions of intensive exploitation, infertility among females was 
higher (72% as against 50%), while among young females, preg- 
nancy was not observed at all. The percentage of pregnant females 
among different geographical populations ranged from 88 to 100% 
in 6 different regions (Zaleker, 1953). 

In Kamchatka from 1952-1962, of 795 females 46.2% were 
pregnant, among second-year females — 27.5%, among third-year 
females — 46.3%, and among older females — 62.5%, and in very 
old females — 20%. Percentage of pregnancy in females ranged (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, 68% of all females 
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 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 vagi- 
nal cornification), the placentation [implantation] of the embryos 
takes place. Their subsequent development proceeds rapidly for 
27-28 days (Kler, 1941). The phenomena of the so-called false 
heat are connected with the period of placentation. The reasons for 
this are 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 273 days. The duration of the latent pe- 
riod of pregnancy in sable is determined to a minor extent by two 
factors: response to [passage of] time and the light factor — change 
in length of daylight. The artificial lengthening of the latter caused 
shortening of the latent period in 50% of experimental females 
(Belyaev, 1950; Belyaev, Perel'dik and Portnova, 1951). 

Parturition takes place 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 in the litter is very tightly linked to the 
degree of favorableness of external environment (nutrition in par- 
ticular); in Siberia [M. zibellina] and North America [M. 
americana], it varies from 1 to 7. In the Pushkin state fur farm, the 
average number of young in a litter equals 3 (Starkov, 1947); in 
the Moscow Zoo, it averages 2.8 (1.5)*. Sable in nature were 
determined to have an average litter size of 2.5 (Kosmakov, 1936); 
546 in Barguzin sable, with the calculation of mortality before the be- 
ginning of harvest — 0.7 (Z.F. Svatosh); in Kondo-Sos'vinsk sa- 
ble — 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 vari- 
able. In individual geographical populations, the average number 
of corpora lutea ranged from 2.3 (Barguzin and Gornoaltaisk sa- 
bles) 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 de- 
pending on the general stability of sable food; thus in the region 
of Barguzin preserve, with little food, the average number of 
corpora lutea was 2.3 (1952), and with an increase in food (1953), 

*Meaning of number in parentheses not clear — Sci. Ed. 



815 



it increased to 3.2 (Baevskii, 1956). In the Ivdel'sk region, in 
1951/52 this indicator was 4.0 and in 1952/53 — 4.5; in Tyumen 
oblast — 3.1 and 5.0 respectively (Zaleker, 1959). 

Variations in the average number of corpora lutea with age 
were also noted in sables. In young female Barguzin sable in 1952, 
it was 2.7 and in 1953 — 2.3; in adult females in 1952 — 3.0, and in 
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 
corpora lutea per female, and in all females of this age — 1.08: 
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 
number of corpora lutea in all very old females was 0.6. For the 
entire period, the average number of corpora lutea per pregnant 
female was 3.96, from 1 to 9. Calculating all females, the average 
number of corpora lutea was found to be only 1.93 (0.51-3.08). In 
second-year females, corpora lutea were (from a total of 1033) 
21%; in third-year females — 30%; and in older females — 49%. 
Females 2-7-year old constituting 22% (on average) of the popu- 
lation, give 50% of the potential offspring, but the second half by 
older females, which constitute only 11%; i.e., the latter are poten- 
tially more (by 2 times) fertile. A definite dependence was estab- 
lished of potential fertility on the nutritional level of the female, its 
weight, and helminth infections (A.A. Vershinin). 

Data on real fertility of sables in nature are extremely in- 
adequate for introducing corrections to potential fertility; however, 
it 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 the number of embryos even discounting embryonic 
mortality. 

Growth, development, and molt. Young sables are born help- 
less, blind, and with membrane-covered ear openings. At that time, 
they are covered with sparse light-gray down. 



816 

The average weight of newborn sables is about 30 g with body 
length of 11 to 12 cm (Starkov, 1947). A single pup may attain 34 
g, while in large litters, it is reduced to 25 g. 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 start to leave the nest at the age of 45 days. 
Lactation ends when cubs have attained 2 months of age. The 
utilization of animal 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, 
at two months — 600 g (Starkov, 1947). By July (the breeding pe- 
riod of the parents), they already weigh about 800 g with a body 
length of 39-40 cm, nearly attaining the dimensions of adult sa- 
bles. With the transition to independent feeding, the growth of 
sables noticeably increases. 

Among 195 newborn sable cubs, 49.2% were males and fe- 
males — 50.8%. Investigation of 1139 individuals from 1918-1930 
showed that 56% were males and females — 44% (Belyshev, 1950). 
According to other data (2,492), males in the population consti- 
tuted 54.4%, and females — 42.6%. Differences have been observed 
by regions (Nadeev and Timofeev, 1955). In Kamchatka, during 
1939-1962, 3,967 individuals were investigated — 52.9% males were 
found, females — 47.1%. In the same place, according to track 
counts, from 1950-1958, 54.2% males were found and females — 
45.8% (A.A. Vershinin). In both cases it is possible that the per- 
centage of males may be overestimated, in the first 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 
sables have few enemies among other animals. In this respect, 
young sables 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 
considered as a factor affecting population numbers. 

In nature, epizootic diseases of sable are undoubtedly present, 
but this question has been poorly studied. A 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 
were favorable for the development of epizootics. Among diseases 
that have been established are encephalitis, coccidiosis and skin 
fungal diseases. 

In the sables, 14 helminthic species have been noted (Petrov, 1941). 
However, their pathogenicity is, in the majority of cases, unknown. 

Mortality of sables is quite significant. Its magnitude is deter- 
mined by many causes. First of all, the mortality of the population 
and its 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 to 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 popu- 
lation age groups were of the following: up to one year — 42.6%; 
1-2 years— 28.5%; 2-3 years— 23.8%; above 3 years— 4.7%. Thus, 
in spite of entirely favorable ecological conditions, the 
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 — 37%, 
moderate age — 37%; old — 26%; females were 59.4%, 31.2% and 
9.4% respectively (Favorskii, 1935). Mortality among females was 
found to be significantly higher than that among males. 
548 In 1952, in the commercial harvest on the territory of Barguzin 

preserve, young females constituted 33%, adults — 50% and old, — 
16%. In the following year, young [females] were 49.2%, adults — 
42%, and old — 6.8%. This appeared to be the result of intensive 
harvesting. In that same year, in an area where commercial har- 
vesting had long been intensive, the percentage of young was 60.7%, 
of adults — 39.3% and old females did not appear at all (Baevskii, 
1956). These circumstances make it possible to estimate the inten- 
sity of exploitation through the age composition (Table 53; Nadeev 
and Timofeev, 1955). 

From all these data it is evident that the size of annual growth 
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 


Vitim 


Angara 


Sayans 


Kuzbass 


Altai 


Average 


Less than 1 year 
About 2 years 
Older than 2 years 
Particularly old 


17.6% 
30.9 
26.7 
24.8 


32.0% 
20.8 
18.2 
29.0 


25.1% 
25.1 
24.3 
24.5 


14.5% 
21.5 
17.2 
47.0 


10.5% 
12.5 
14.6 
62.4 


28.5% 
27.1 
21.3 
23.1 


Examples studied 


165 


131 


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 
to 36.8% (average; 31.6%). In northwestern Yakutiya, yearlings 
comprised 1/3 of the population, and in the Oleneksk region 
(1950/51), 37% (Tavrovskii, 1958). 

Age analysis of 1762 sable skulls from Kamchatka (collected 
1945-1962) gave the following results (Table 54). 

The decrease in the percent of young females is explained by 
their great utilization; there is information about the increased 
mortality 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): 

% % 

1952/53—61.5 1957/58—14.3 

1953/54—14.9 1958/59—81.5 

1954/55_51.9 1959/60—43.7 

1955/56—41.2 1960/61—15.0 

1956/57—28.6 1961/62—50.5 
For 10 years, the average percent constituted 40.3 (14.3-81.5%). 

549 In Kamchatka, for the same decade the size of the potential 
increase was obtained based on the percentage of adult females 

Table 54. Age composition of Kamchatka sables from 1945-1962 



Age 


Total 


Males 


Females 


Juveniles 


27.3% 


21.5% 


23.0% 


18-22 months 


22.1 


19.8 


24.3 


In the 3rd year 


19.4 


19.3 


19.5 


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 selec- 
tion of harvested animals as regards their age (A. A. Vershinin). In 
Kamchatka, 32.5% die during the embryonic and early postembry- 
onic periods, and in the unfavorable years — 50% of the potential 
increase. 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 competi- 
tors. 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 ani- 
mals of the taiga are considered competitors of sable in hunting 
mouse-like rodents. Siberian weasel, ermine and weasel are them- 
selves 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 




% 


of 


corpora lutea 


growth 


the harvest, % 


1952/53 


30.0 




1.36 


40.8 


61.5 


1953/54 


43.5 




0.84 


36.6 


14.6 


1954/55 


32.5 




2.46 


80.0 


51.9 


1955/56 


27.4 




1.80 


48.5 


41.2 


1956/57 


36.0 




0.79 


28.5 


28.6 


1957/58 


43.5 




3.08 


134.0 


14.3 


1958/59 


22.7 




2.01 


45.6 


81.5 


1959/60 


30.4 




0.51 


15.5 


43.7 


1960/61 


41.3 




2.34 


96.6 


15.0 


1961/62 


25.3 




2.70 


68.4 


50.5 


Average 


33.0 




1.86 


61.3 


40.3 



820 

Population dynamics. Changes in their numbers is a character- 
istic of sables. Changes brought about by deep reconstruction of 
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 
infrequently, and not everywhere. In cases when its density in- 
creases, the possibility of contact grows, and serious epizootics 
begin to appear among sables. Where human activity is excluded 
or weak (preserves), fluctuations in sable number are insignificant 
550 and are determined in 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 
are repeated, on average, every 10 years, with deviations in the 
range of 9-11 years. 

In Kamchatka, voles are of decisive importance as sable food 
(see above). Their mass reproduction there has a three-year cycle; 
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 fertility of young females. With intensive repro- 
duction when the main food is plentiful, the sable population at- 
tains a younger average age and therefore, in the following year, 
breeding intensivity is reduced. In the third year, the age structure 
becomes even less favorable for productivity of the population, 
and it 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 popu- 
lation is determined by the availability of the main food: 1) environ- 
mental factors which determine changes in abundance of voles, and 2) 
by changes in population structure which change the level of fertility. 

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 
fluctuation attains an interval of 6 [years], in such regions with a 
prolonged cycle, it is 2-3 times lower; i.e. the population is more stable. 

In the Glacier National Park (Montana, USA), the process of 
numerical change in concrete local populations in an area of 1,554 
ha was studied by means of tagging and recapturing animals (1952- 
54). 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 insuffi- 
cient 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 itself may 
be met with rarely and accidentally; it is easier to succeed with the 
help of a good laika dog. The sable, driven up a tree, somewhat 
resembles a small cat, 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 
head and somewhat shorter tail. The male can be easily differen- 
tiated 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 often find sable scat on taiga paths, on stumps 
and fallen logs. It is sausage-like in form with blunt ends, about 1.5 
cm in diameter, and a length of about 6-7.5 cm. In it are usually found 
hair and small bones of voles, shells of nut pine "nutlets", fruit and 
insect remains. The scat gives off a musky odor. 
551 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 sum- 
mer. 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 
depression of dimensions 10 x 25 cm. On packed snow, sable 
tracks 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. 

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 each other, as in sable, but somewhat overlap each other 
at the heels; this is less marked in the tracks of pine marten and 
kidas. When jumping from a tree, the pine marten does not leave 
the tail print on the snow, but the sable does. Passing under the 
crown of a tree, the sable greatly shortens its 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 settled and when the animal sinks in not 
more than 3-5 cm. 

Usually, the sable does not run the whole day with one gait. 
The length of the leap depends upon snow density. In soft snow, 
leaps are shorter and the sable places 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). 

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 
paired prints of a track in females average 11.0 cm (9.4-11.9), of 
a male— 13.3 cm (12.0-14.6). 

In the most typical gait, which make paired prints, the poste- 
rior pair of feet lie in the prints of the anterior pair. One of the feet 
is usually displaced forward, and there is no regular alternation of 
the right and left feet. A triple print, in which one of the posterior 
feet does not lie in the print of the anterior foot, is infrequently 
observed and is usually noticed when there is little snow, when the 
sable feels firm support. It rarely places all of its feet separately 
(as in red fox). 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 is the most valuable fur-bearing animal in the USSR 
although, as a commodity, the quality of the skins is not of the 
same value in different geographical regions and even in the same 
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 reintro- 
duction of sable were undertaken in regions where it had been 



824 

extirpated. As a result, the number of sable and the area inhabited 
by it increased considerably. The 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 
33% of the sables caught each year come from Krasnoyarsk Terri- 
tory, about 20% — from Irkutsk district, about 16% — from 
Khabarovsk Territory. With all of this, at the present time, not all 
the areas in the Siberian taiga suitable for their occupation have 
been taken up by sables. At the present time, the tendency is to- 
wards increase in the number of light-colored sables. Therefore, 
there stands the problem of devaluation in the direction of natural 
selection of sable with the entire improvement in the quality of the 
skins on a large scale. 

A very important measure for management of the sable popu- 
lation is to determine correctly a guide to the amount of utilization 
(the percentage of the admissible catch) and maintain it. For a very 
high density, the catch should be accelerated. The example of 
Kamchatka can illustrate this situation: a catch of 7.5-8.0 thou- 
sand sables in a season does not lead to a decrease in their number, 
but a catch of even 8.5-9.0 gives this effect. In Kamchatka, it is 
admissible to take 27% of the autumn pre-harvest number (A. A. 
Vershinin). In other places— 25% and 30% (Baturin, 1930; 
Vershinin and Dolgorukov, 1947). It 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 deadfalls, snares, and live-traps of the box 
type are also used. The hunting of sable by baits helps to obtain 
them alive, which is 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: 
1) strict maintenance of rules and norms of licenced harvest; 2) 
prohibition of selective destruction (which is possible by baits of 
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 and shooting of only the lowest qual- 
ity sables ("mekhovaya" and in some cases, "vorotovaya"). 

The greatest efforts should be directed towards the develop- 

553 ment 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 more convenient and 

easier to develop (P.Yu.). 

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., 2, p. 1029. Sub- 
stitute for martes. 

1820. Mustela sylvatica. Nilsson. Scand. Fauna. Dagg. Djur., 1, p. 
41. Renaming of martes. 

1827. Martes vulgaris. Griffith. Cuvier's Anim. Kingd., 5, p. 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 district. 

1926. Martes martes ruthena. Ognev. Ibidem, p. 49. Dmitrovsk, 
Moscow province. 

1941. M.(artes) m.{artes) ruthena borealis. Kusnetzov. Tr. Mosk. 
Zootekhn. in-ta, 1, p. 126. Shenkursk region, Arkhangel'sk 
district. Nomen 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. zapov., 5, p. 179. Pechoro-Ilychsk preserve, 

upper Pechora (V.H.). 

Diagnosis 

Tail length with terminal hairs more than half that of body. Color 
monotone, sharply outlined patch of various forms on throat and 
chest, usually varying intensities of yellow color, sometimes white 
or almost white, and as a rule, not forming two strips extending 
backwards to base of forelegs. Head not lighter than back. Bony 
auditory bulla relatively shorter and more widely separated than in 
sable. Inner half of upper molar considerably larger than outer 
(V.H.). 

Description^^ 

In its general appearance, the pine marten in winter pelage is very 
similar to the sable, but looks somewhat larger, coarser 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 
stretched hind legs usually for more than a quarter of its length. In 
contrast to the sable, the terminal hairs of the tail do not form a 
blunt, but a tapering tail tip. This tip is sometimes strongly elon- 
gated (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 ter- 
minal 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 
554 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, how- 
ever, 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 long- 
tailed, with less broad feet and not so big-eared. In its manner of 
standing and moving the marten is entirely similar to sable, but 
differs considerably from the stone marten, which mostly "crawls" 
and rarely proceeds by jumping. This is connected, in part, by the 
height of the animal at its shoulders: in the pine marten, this consti- 
tutes about 15 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 

The pelage of marten is similar in structure to that of sable, 
but is somewhat rougher and less dense and fluffy. Nevertheless, 
it 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 
555 guard hair. Length of the guard hairs is 23-27 mm, with a thick- 
ness 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 mm and 14.5-15 microns (Pavlova, 1951). Thus, the win- 
ter hairs although longer, are relatively much thinner. In the sum- 
mer coat of marten, the sole and finger cushions as well as the 
claws are easily seen. In winter, the tail is 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. 

A 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 exten- 
sion is absent) or greatly reduced. In some instances, it is divided 
into separate parts and, rarely, may be reduced into 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 it is 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). 

Though the general color type is retained, its tone and 
intensity undergo considerable individual fluctuation. One may dis- 
tinguish four main color types of martens which are accepted in 
the fur industry (furriers terminology). 

Dark blue — fur dark-chestnut without reddish tints, underfur 
bluish-gray at the base and light-gray at the tip. 

Blue — pelage lighter, chestnut colored, underfur gray. 

Dark sandy — fur brownish ("dark-sandy") with reddish tints, 
more distinct on the sides. Underfur gray at base and light-sandy 
color at hair tips. 

Sandy — pelage light-brownish ("sandy") with light-yellow tint. 
Underfur gray at 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 in sable and the phenomenon of parallelism 
itself is less marked. 

The color of the throat patch is, as a rule, yellow of varying 

intensity, changing from slightly ocherous to bright intensive 

ocherous, almost orange. Near-white and white patches occur. There 

556 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; north- 
ward, intensity of its color decreases, and it lightens. The same 
picture is 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 
are sometimes found, or having a form, similar to the form typical 
for stone marten. Such individuals are very rare, and only one 
among several hundreds are met with — in the Urals, one among 
900 (Pavlinin, 1959)^8. 

The density, luxuriance and fineness of the pelage i.e. fur qual- 
ity, undergo marked geographical changes which are, however, 
much less than in sable (see section "Geographic variation"). In- 
dividual and geographic variation in marten are on the whole, much 
less than in sable. Geographical localization of types of individual 
variation so characteristic for sable is 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, which is much darker 
than in winter. The tail, particularly its tip, and the feet are darker 
than the trunk — blackish-brown. The throat patch is usually brighter 
than that in winter marten. Individual variation in color of summer 
fur of martens is much less than in winter. 

There are no sexual and age differences in color and character 
of the pelage (fur quality). Young in the first autumn put on a coat 
that is not distinguishable from the winter coat of adults. 

The skull of pine marten is very similar to the skull of sable. 
It differs from it by the not so elongated brain case and by its 
shorter and more widely separated bony auditory bullae — the dis- 
tance 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 
dental formula is as in sable, but the inner blade of the upper 
molar is usually somewhat larger and the difference between the 
inner and outer blades is 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, in the somewhat 
lighter structure of the skull. In the martens of Volga-Kama basin, 
the condylobasal length of the male skull is more than 78 mm and 
the zygomatic width is more than 45 mm; in females, 
correspondingly — less than 76 mm and less than 42 mm (Aspisov, 
1959). Age changes in the skull are significant and, in general, 
correspond morphologically to those described for sable (age con- 
formity is not established). 

The skeleton of the tail is composed of 15-22 vertebrae, i.e. 
on average, several more than in sable; Caucasian martens have 
18-21 caudal vertebrae; Pechora— 17-20 (17—5%, 18—40%, 19— 
25%, 20 — 30%; Yurgenson, 1947). Male martens of the middle 
558 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). The clavi- 
cles, which are rudimentary in all species of the genus, are articu- 
lated 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 more or less with 
sable, usually averaging somewhat less than the latter. Within the 
species, body length is 380-580 mm, tail length, 170-260 mm, 
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 
differ in the following manner: body length of males — 422-439; 
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; weight of 
males, 795-850 gm, females, 542-620 gm^^ Weight of the female 
skin constitutes 68.4% of the weight of the male skin. 

"Weight of carcass given without skin, i.e. it is actually somewhat larger. The 
weight of a freshly skinned hide in the same region comprised from 96 to 159 gm 
(Pavlinin, 1959). It is possible that some other measurements provided by the author 
(body length?), are also not standard, but are of the same type and give a represen- 
tation of the relative dimensions of the different sexes. 



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 

Body length (mm) 93 405—473 102 365—420 

Tail length (mm) 104 185—228 100 170—210 

Hindfoot length (mm) 84 78—93 78 70—84 

Ear height (mm) 6 45—47 4 43—46 

Weight" (gm) 59 670—1050 49 484—850 



Condylobasal length of male skull, 72.1-88.0 mm; females, 
71.1-83.0 mm; zygomatic width of males, 41.4-53.8 mm; females, 
38.2-49.2 mm; skull height in males, 27.4-34.9 mm; females, 27.0- 
31.8 mm (from material of Kuznetsov, 1941). Weight is 580-1800 
gm. 

In adult martens (older than 2 years), length of the os penis is 
38.9-45.7 mm, M 42.3 mm; weight, 0.20 to 0.35 gm, M 0.29 gm. 
In young, length is 35.1-38.3 mm, M 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 may be from 4.3 to 11.5. Weight of the liver of males 
is 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 Kidas.3' In those places where sable and marten occur together 
and particularly in the Urals, in the 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 estab- 
lished a restriction on kidas fertility — females are fertile in back- 
crossing both with sable and with marten; in this combination, 
males were shown to be infertile. Fertility of kidas among 

'"The given figures, some at least, were apparently taken from frozen carcasses 
and assigned to the weight of the body without hide. See the data on other internal 
features in the present work. 

"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, suffi- 
ciently extensive experiments of this kind. 

This complex hybridization mentioned above leads to a 
situation that on one level individuals with more or less defined 
intermediate characteristics more similar to martens — "marten-like" 
type, and also those in 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 char- 
acterization 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 dif- 
fering 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. 

Kidas of the same reserve are also of different size; on the 
average they are smaller than sable (which is larger than marten 
in the Pechora), but larger than marten. Individual animals are 
larger than the largest martens and are nearly of the same size as 
the largest sable. ^^ The dimensions of kidas (24 males, 10 females) 
560 are as follows: body length of males — 390-M 452-485 mm, fe- 
males— 365-M 406-450 mm; tail length of males— 170-M 187- 
240 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 
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- 
tion between sable and marten. In sable of all ages, this is equal 
to a mean 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 
width 52.1-M 55.6-58.1 mm; skull height 36.1-M 38.5-42.8 mm 
(10 spec). 

The structure of the os penis in the majority of cases (more 
than 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, 35% have an intermediate characteristic between martens 
and 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 distribution of different upper molar structures among 
individuals is as follows: 41.6% of kidas have "sable-like" teeth, 
41.6% — "marten-like" type, and 16.8% — intermediate characters. 

559 "According to other data (Yazan, 1962), kidas of Pechora-Ilych reserve (14) are 

larger than both martens (463), 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 are not clear (fresh carcasses with fur, frozen carcasses 
with fur, or frozen skinned carcasses?). 



836 

A marten-like postorbital constriction is present in 8.3% of kidas, 
sable-like — 33.3%, and transitional type — 58.4%. It 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, tail as in pine marten; in craniological 
features, more sable characters, etc. Clear intermediate forms are 
sharply distinguished, such as light Ural sable with long marten- 
like tail. 

In different places, evidently in connection with the correla- 
tion 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 — 
for example, on the Pechora: marten — 58.8%, kidas — 29.6% and 
sable — 11.6% or 80%, 7%, 13% respectively. When marten and 
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 
even 65 and 35%" (V.H.). 

561 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 

This constitutes about half of the species range. It is 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 
appeared ("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, 
the formation of races of, hybrid origin, etc.) are also rejected. 

At the same time, the author assumes that kidas in nature all "appear under 
exceptional conditions" (page 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 little to 
the north (the most northern point of occurrence in the 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 
coast, to which the range almost everywhere, extends except, ap- 
parently, 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. 
Parovshchikov). 

Beyond this line long transgressions occur — the Kola Penin- 
sula to lokan'ga on the [White] Sea coast, 50 km from the forest 
border (Smirnov, 1901) and at Kiya station, on the western shore 
of the Kanin, approximately 200 km north of the place of perma- 
nent 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 cross- 
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 the Ob', the range occupied the region of the lower Nazym 
where it 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 this parallel, in general, or a little farther to the north, 
it 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 south- 
west to the sources of the Dem'yanka (Skalon and Raevskii, 1940). 
Thence, moving in the same direction, it crossed Irtysh and the 
lower Ishim and, proceeding a little northward to 56° N. lat., through 
Golyshmanov and Yalutorovska regions, it passed on to the Urals 
through Sysert' to Nyazepetrosvk. Thence, it directed itself south- 
ward to include the Ural [river] extending into its valley to reach 
Chkalov. 

An isolated section of occurrence, analogous to those found in 
the 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. It has as its base 
the Urals between approximately 65" N. lat. and 56° N. lat., 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 in the 
Shuryshkar region, at approximately 66° N. lat., i.e., only a bit 
more to the south than on the right bank of the Pechora. Appar- 
ently, at this latitude, it also crosses the Ural [mountains]. 

Farther, the border descends to the neighborhood of Berezov 
on the Ob' and, going eastward, encompasses the basin of the 
Kazym — the Khetu and Sorum rivers, the Trom-Yogan basin (Trom- 
Agan, 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 

*Not clear; the range as described and mapped does reach the middle Ob' ; unless 
this is meant to refer to 1920-30 period (see Fig. 211) — Sci. Ed. 







■v^ 03 3 



500 



^ V p 



840 




563 Fig. 211. Borders of the distribution of pine marten, Martes {Martes) martes L. in 
western Siberia (V.G. Heptner): 1— Border at the beginning of the 50's of the XX 
cent.; 2— Same in the 20' s and 30' s of the XX cent.; 3— Proposed limits of the range 

in the XVII cent.; 4 — Transgression south of Petropavlovsk. 

of the Vasyugan. Thence, the border of the range passes to the 

564 Irtysh at the mouth of the Тага 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 со.; Ornev, 1931) and Savransk for- 
est (west of Pervomaisk), passing toward Dnepropetrovsk (Samarsk 
pine forest, the Samara river which flows into the Dnepr from the 
left; Yurgenson, 1932) and farther, to Izyum and to the mouth of 
the Oskol in Donets (former Izyumskii Co.; 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 east- 
ward, reaching the Volga at Saratov or a little below (it occupies 
the forest-steppe on the left bank; El'pat'evskii, Larina and 
Golikova, 1950). 

In the above-described segment of the southern border of the 
range, the marten extends into the forest-steppe zone, and its ac- 
tual outlines are very complicated. The described line passes through 
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 
number of places, over the unforested steppes of interfluves. 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. 



842 

From Tambov or from the sources of the Don, in an unclear way, 
it descends to Voronezh and then along the Don 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 allot- 
ment) and neighboring forest massifs, in the forests along the 
Bityug, and particularly around Bobrov and in the Khrenovsk 
forest (N. Severtsov, 1855; V.G. Heptner), around Novyi Kurlak 
and in several other places, in the Kamennaya Steppe (Dokuchaevsk 
Experimental station around Talovaya station), and in the Shipov 
forest near Pavlovsk (Ognev and Vorob'ev, 1923). 

Beyond the Volga the border, apparently starting near the city 
of Engel's, goes northward along the floodplain of the Volga to 
the mouth of the Bolshoi Irgiz. From there, it passes along flooded 
forests of this river eastward nearly to Pugachev, then returns west- 
wards to the Volga and extends along its right bank to the mouth 
of the Malyi Irgiz, rising somewhat to the east in the extreme 
lower reaches of this river. Thence, the border line again directed 
itself northwards, along the Volga floodplain, reaching the bend of 
the Samara; apparently, the southern border passes eastward along 
the Samara river (N.I. Larina) towards the Ural and joins the above- 
described border, which passes from Siberia to the Ural [river]. 
Along the bottom land of the Ural and the rivers flowing into it 
(Burli and others), the marten goes down the river to Ural'sk and 
descends a little below it (transients known to Gur'ev; Sludskii, 
565 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 the European-Siberian part of the range 
throughout its whole extent from the Barents Sea to the Black Sea 
is formed by the state frontier. 

The Caucasian portion of the marten's range occupies the for- 
est regions of the Main Caucasus range northwards, including the 
foothill forests. The range apparently also includes the forests of 
the Terek valley lying on the plain. Information on the marten in 
the Parabochevsk forest near Shelkovsk station and in the mixed 
forest tracts of the Terek valley (Heptner and Formozov, 1941) 
relate, evidently, to this species. In the west, the range includes the 
basin of the Pshekha river (left branch of the Belaya), and perhaps 



843 

extend somewhat farther. To the east, the range reaches the inte- 
rior forests of Dagestan in the basin of the Avarsk Koisu (Dinnik, 
1911). Marten also occurs, apparently, in the foothill forests south 
of Makhachkala, but positive information on this is 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 Cauca- 
sus 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 Adzhar- 
Imeretinsk and Trialetsk ranges from Borzhomi and Bakurian' to 
the Belyi Klyuch. To the east of Akstafa-Chai and Debeda-Chai, 
i.e. the meridian centering on Lake Sevan, this marten is absent or 
extremely rare. 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, particu- 
larly at Talysh (Radde, 1886; Satunin, 1896) were denied by all 
subsequent authors. However, the latest data 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 
south, the more it becomes sporadic, in accordance with the rela- 
tive area and 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 con- 
ditions its distribution is becoming all the more sporadic in the 
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, it was not, and is 
not now subjected to evident changes. The described range is the 
present species range, in all events in the European part of the 
Union and the Urals. 

As for the Trans-Ural and western Siberia, the history of the 
marten's range is, apparently, more complicated. In the literature, 
there are quite a few assertions that earlier the marten was not 
distributed beyond the Urals and 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). Concern- 
ing 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 phenomenon is usually considered to 
be the sharp reduction in the number of sables and its extermina- 
tion in several districts. With this, a suitable ecological niche for 
566 the marten was freed-up. With a normal number of sable, the marten 
could not penetrate into the east because of its "resistance". It has 
been considered that the cause might be change in natural 
conditions — 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 West Siberia, the marten was 
already known a long time ago. Thus, in the 17th cent., it lived 
here along the border of the forest zone and throughout the forest- 
steppe in quite considerable numbers and still remained in the 18th 
cent., though in very small numbers. Pallas also took note of the 
marten near Тага. The marten existed not only in Tobolsk and Тага 
counties of that time, 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 Surgutsk 
county (1630, 1650, 1660) or it was met with here very rarely 
(Kirikov, 1960). Evidently, marten was distributed in southern Si- 
beria far to the east, but in the north, it, apparently, did not reach 
the [Trans-Ural] plain. 

Apparently during the 18th and 19th cent., marten (as well as 
sables) completely or almost completely disappeared in a signifi- 
cant part of its previous West Siberian range. It is difficult to 
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. 

On the basis of all these referenced materials, the history of 
the marten range in western Siberia more correctly presents the 
following aspect: in southern regions — along the southern part of 
the forest zone, the marten lived in the forest-steppe 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 
strongly, it disappeared or almost disappeared in western Siberia. 
In recent times, the range is being re-established, especially inten- 
sively in the last decade. However, the full range in the south is 
still not re-established. In northern Siberia, the marten is settling 
in new regions, where, previously, it was absent. There, coloniza- 
tion has been taking place, in our day, especially intensively in the 
30' s. Therefore, externally similar phenomena have fundamental 
differences. 

The reasons for these changes in range are not understood in 
detail and require careful investigation; evidently, both of the above- 
mentioned phenomena — biocoenotic (relationship between sable and 
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 numbers were, probably, more signifi- 
cant; in the north, equally with the biocoenotic factor, change in 
the character of forests 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 exploi- 
tation, 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 Iraq, and to Iran in two extensions — along the Elburz 
system, it 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 its occurrence in northeastern China, where 
the pine marten is supposedly even more common than sable, and 
is 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 
applies, of course, to the true marten (see in particular Lukashkin 
and Zhernakov, 1934). References to the occurrence of marten "in 
Siberia, Turkestan to Manchuria" are also improbable (Brass, 1926) 
(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, estab- 
lished 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" sub- 
species, and several are a priori doubtful. The characteristics of 
marten skins as a commodity fur, apparently, served as the stimu- 
lus for distinguishing some forms. 

A new revision of geographic variation in the species is nec- 
essary. The scheme of B.A. Kuznetsov (1941, 1952) is presented 
below with a few modifications, critical remarks and nomenclatorial 

568 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, sylvestris, abietum). 

Size large, tail long. Skull large and massive. 

Fur on spine beautiful dark-chestnut color; guard hairs dark- 
brown, underfur bluish-gray, tips lighter and paler. On sides, fur 
slightly lighter. Withers and upper part of neck and head some- 
what darker than spine. Muzzle brownish, slightly lighter 
posteriorly. Throat patch usually bright yellow. Belly dark brown- 
ish, darker than sides of body. Base of tail same color as back, to 
brownish toward its tip. Feet dark-brown. Pelage fluffy and fine. 

Body length of adult males 480-530 and to 550 mm, of fe- 
males 400-450 mm; tail length of males 250-280 mm, females 
230-260 mm (Schmidt, 1943; Middle Europe). 

Condylobasal length of male skull (22) 81.0-M 86.26 ± 0.46- 
88.0 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, 



848 

М 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 north- 
ern Kareliya. 

Outside the USSR — western Europe, except the Mediterranean 
region. 

The identity of Kola martens with our western and southwest- 
ern ones is 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, 
underfur light bluish-gray with reddish-tawny hair tips. Sides lighter 
and grayer than back, underfur on the sides also such color as 
back, but with pale hair tips. Belly darker than sides, tail at base 
colored like back; at end reddish-brown. Distal part of legs dark- 
tawny. 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 
mm), length of tail 195-M 218-260 mm (Yurgenson, 1947). 

Condylobasal length of male skull (32) 75.3-M 80.19 ± 0.42- 
84.8 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- 
M 42.22 ± 0.32-44.3 mm; skull height of males 27.4-M 31.07 ± 
0.14-33.3 mm, females 27.0-M 28.84 ± 0.28-31.1 mm. 

In 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 line. 

Absent outside the USSR. 

3. 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 
hair tips. Tail tawny, lighter at basal part and darker at tip. Legs 
dark-tawny. Throat patch light-creamy or pale-yellow, rarely pure 
white. Pelage fluffy and soft. 



849 

569 Dimensions similar to preceding form. Body length of martens 
of upper Pechora (67) 380-M 433-460 mm; tail length 170-M 
198-235 mm. Weight (42) in winter 580-M 784-1090 gm 
(Yurgenson, 1947). Weight of 100 skins 8.5 kg. 

In European part of the USSR north of the Vologda-Gor'kii- 
Kazan line, 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 grayish- 
tawny, 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 
as back, its tip dark-tawny. Fur fluffy and soft. 

Condylobasal length of male skull (7) 79.8-M 83.57 ± 0.78- 
87.0 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- 
M 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 by reddish tone on the 
posterior part of the back which are not developed in the other two 
forms (B.A. Kuznetsov). 

5. 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 
tawny, undercoat bluish-gray, moderately dark, hair tips sandy- 
yellow. Sides lighter than back with a grayish tint. 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 tail dark 
chestnut, tip brownish. 

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 
mm (85.3 mm)35-53.0,* females (14) is 74.6-M 75.43 ± 0.45 mm 
(78.3 mm)-81.2; zygomatic width of males 43.1, M 47.42 ± 0.45 
mm (50.9 mm)-53.0, females 41.2— M 44.14 ± 0.50 (43.7)-48.3 
mm; skull height of males 28.4— M 31.87 ± 0.33 (32.1 mm) 
-34.0 mm, females 28.1-M 29.93 ± 0.31 (29.5)-31.8 mm. Weight 
of 100 skins 11 kg. 

In Middle Caucasus and Trans-Caucasus. 

Outside the USSR — in contiguous parts of Turkey and Iran. 

A 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 — South Italy south of Abruzzi (V.H.). 

570 Biology 

Population. The pine marten, restricted in its distribution to the 
forests of the European part of the USSR, is relatively few in 
numbers. The entire size of the population of this species before 
the October Revolution was approximately 40 thousand individu- 
als or a little more. The number of pine marten greatly increased 
after the October Revolution. Very approximately, its numbers may 
now be considered equal to 200-300 thousand. Some estimations 
of numbers of the pine marten refer to tannery figures for the 
periods of irregular exploitation (1922-1925). The Northern Ter- 
ritory produced about 21% of the annual catch of marten skins; at 
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 in error; too 
small. M may be correct — Sci. Ed. 



851 



The forest marten is, within its range boundaries irregularly 
distributed, a fact conditioned by the percentage of forested areas 
in the different regions and the degree of suitability of habitat 
(nutritional conditions, availability of prey items, nest and refuge 
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 densities. 

Habitat. In all features of its structure and ecology, the pine 
marten is closely associated with forest. Within such forests, it 
reveals very significant plasticity in relation to external environ- 
mental conditions. It is encountered from the sparse pine forests of 
the Kola Peninsula and northern taiga to the oak forests of Medi- 
terranean type, the "maquis" of Sardinia and tall beech forests of 
Adzhariya and Kolkhida. 

Within the limits of such landscape zones are the most pre- 
ferred habitats of the pine marten. In the coniferous forest zone, it 
is 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 mar- 
ten were met with twice as often as in pine forests and other 
plantations. A series of daily trail surveys showed that 81.5% of 
the total length of trails are found in the spruce forests, 11.4% — 
in pine forests and only 7.1% — in other plantations. These distri- 
butions changed in the different seasons depending 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 among the 
spruce forests — complex spruce forests, the spruce-aspen forests 
and the so-called pine-spruce aggregations (pines, spruce-moun- 
tain sorrel, and spruce-bilberry). In the zone of the broad-leaved 
forests, it prefers the large broad-leaved forests and their combina- 
tion with montane pine forests, in the montane forests of the north- 
ern Caucasus, the fir-beech and beech forests, and the forests lying 
at their upper limits. Class I 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 II quality areas are oaks, beech, 
pear, chestnut, aspen and alder groves in the broad-leaved [forest] 
zone (Ryabov, 1958). 

Quality (bonitet) of habitat for pine marten regularly increases 
from north to south parallel to the change in forest biogeocoenoses 
and simultaneously with shifts in the nutritional regime. The eco- 
logical optimum of the species lies in the montane forests of the 
571 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 polypha- 
gous 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 

The relative weight 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 geographic situation of the locality which all the more deter- 
mines 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) 
of each food type. 

There is a definite regularity in geographic variation of pine 
marten foods — carnivory of the diet increases from south to north, 
and omnivory — from north to south. This is also reflected in vari- 
ation of the chewing apparatus (Yurgenson, 1951). From north to 
south, mammals and birds, especially tetraonids, as well as bird 
eggs regularly decrease in the 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 — in the snowy period. Its average 
yearly occurrence is similar everywhere and reflects the plant food 
requirement of the animal. 
572 Seasonal variation of food is also well illustrated. For example 
the squirrel mainly occurs in pine marten food in the snowy pe- 
riod, constituting about 44% occurrence or more while in the 
snowless period — not more than 6-8%. The same may be said 
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 met with significantly more rarely. 

There is a marked sexual dimorphism in foods of the pine 
marten. In the food of the weaker females, which are of smaller 
size and weight, hares, capercaillie, black grouse i.e. 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 in foods are pre- 
sented in Tables 57 and 58. In these, 2,751 data points on nutrition 
of pine martens were used: 1) Lapland — 587 (Nasimovich, 1948); 



854 



574 



2) upper Pechora — 593 (E.N. Teplova and P.B. Yurgenson)*; 

3) Vologodsk district— 205 (Gribova, 1958); 4) Zhiguli— 151 (P.B. 
Yurgenson)*; 5) the middle zone — 220 (Grigor'ev and Teplov, 
1939; Yurgenson, 1939); 6) Northern Caucasus — 1300 feces and 
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 
10 years (1937/38-1948/49), the occurrence of basic food types 
changed within the following limits (Table 58). 

In Table 59 only the average occurrence of the different com- 
ponents 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 


Zhiguli 


Northern 
Caucasus 


Mammals 


70.3 




63.5 


48.7 


83.0 


62.3 


Mouse-like rodents 


34.3 




34.9 


24.7 


72.0 


58.3 


Squirrel 


3.1 




26.3 


11.6 


9.3 


— 


Insectivores 


4.4 




6.7 


4.9 


— 


3.0 


Birds 


34.8 




30.6 


21.5 


4.0 


18.0 


Including 


23.0 




15.7 


7.4 


— 


— 


tetraonids 














Bird eggs 


6.9 




5.7 


4.1 


— 


0.6 


Insects 


10.5 




10.7 


7.4 


24.0 


39.5 


Plant food 


23.9 




25.5 


No data 


14.6 


14.8 


Fruits (including 


17.5 




11.6 


No data 


12.8 


14.8 


berries) 














Table 58. Foods of pine martens in 


the upper Pechora over 


10 years 


Food type 






% of occurrence 




Average, % 




M 


inimum 


Maximum 




Voles 




7.8 




29.4 




23.3 


Squirrel 




13.7 




58.1 




36.8 


Tetraonid birds 




4.0 




24.0 




15.1 


Other birds 




0.0 




23.0 




8.3 


Plant food 




5.0 




39.1 




24.5 


Other food 




6.1 




20.9 




11.8 



"No year given — Sci. Ed. 



855 



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856 

In Arkhangel'sk oblast, from 1949-1961, i.e. during 13 years, 
the occurrence of different foods was as follows (Grakov, 1962) 
(Table 60). 

In one of the provinces of Sweden (Heglund, I960)*, the char- 
acter of winter food and its variation (%) within a three-year pe- 
riod were as follows (1956/57-1957/58-1958/59) 

Mouse-like rodents 44.9 (25.0-61.5%) 

Squirrel 39.7 (25.6-60.7%) 
Passerine birds 7.7 (6.5-9.0%) 

Birds (general) 21.8 
Bird eggs 7.7 

Fruits (including berries) 12.8 

The occurrence of squirrel in marten food increased with the 
decrease in mouse-like rodents and vice versa. 

From the tables presented, it is evident that even within the 
boundaries of one zone (in the European northern taiga), consid- 
erable variation in the degree of occurrence of the main food groups 
is observed. In particular, the range of variation in the separate 
food groups is more sharply marked in Arkhangel'sk district than 
in Pechora, because there (in Arkhangel'sk district) the occurrence 
of squirrel and plant food is lower. The latter results from the 
absence of nut pine "nutlets" in winter food. On the Kola Penin- 
sula the differences are even more marked. 

Table 60. Nutrition of the pine marten of Arkhangel'sk oblast during a period of 

13 years 



Food type 








Occurrence, 


, % 


Average (m) 




Minimum 


Maximum 




Voles 






12.5 




75.5 


45.1 


Squirrel 











17.5 


7.1 


White hare 











9.2 


5.0 


Insectivores 











6.1 


2.0 


Birds 











32.5 


21.5 


Including tetraonids 




2.8 




22.8 


13.6 


Bird eggs 











10.5 


3.4 


Frogs 











9.2 


3.5 


Berries 











43.1 


11.2 


Mountain ash 











17.0 


3.2 


*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 concen- 
tration of pine marten 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 
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 is considerably easier for mar- 
tens, their relative weight in the marten's food increases. 

Species composition of mouse-like rodents utilized by martens 
is variable. It depends not only on their species composition and 
relative abundance in the local forest fauna, but also on their rela- 
tive availability: the more active wood mouse [Apodemus] is al- 
575 ways less common. The forests of the Black Sea coast of the 
Caucasus constitute an exception where, in some places, other 
mouse-like rodents are absent. 

In the past, there was a 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, 
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 food sharply decreases. The destruction of young squirrels in 
the nest 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 
years when squirrels are very few and martens are, on the other 
hand, abundant, these carnivores may destroy up to 30-35% of all 
squirrels; in other years, the importance of squirrels is consider- 
ably lower. It is believed that in the European North (Grakov, 
1962) the pine marten lacks the power to affect the dynamics of 
squirrel numbers and cannot determine their number. Marked losses 
in local populations of squirrels are only caused by marten in those 
rare years when the low number of squirrels is accompanied by a 
low number of the tetraonid birds and mouse-like rodents, while 
martens are, at the same time, numerous. 

An abundance of squirrels in nature does not cause an increase 
in their occurrence in marten food, especially if there are many 
voles. As was noted in Pechora, increased occurrence of squirrel 
in marten food occurs in connection with the increased occurrence 
in nature of weak and sick squirrels. 

In the nutrition of the pine marten of the European taiga and 
some parts of the mixed forest zone, tetraonid birds acquire an 
essentail importance, especially capercaillie [Tetrao], black grouse 
[Lyrurus 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 mar- 
ten mainly in their snow burrows and night resting places and, 
therefore in summer, their occurrence in marten food decreases 
significantly and becomes more or less accidental. By means of 
tracks in the snow, it is frequently possible to trace the unsuccess- 
ful 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). White hare also occurs only in 
the food of males. 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. 

The capture of small birds is 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 in certain 
winters and in fairly great numbers. This is connected with the 
death of moles through strong freezing of the taiga soil in years 
with little snow (Teplova, 1947). The eggs of tetraonid birds, mainly 
those of capercaillie, in winter food of marten is also due to spe- 
576 cial conditions. This is connected 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-Tyan- 
Shanskii, 1959). 

Sometimes, remains of frogs and their eggs are found among 
the remains of marten food in winter. This is observed in those 
winters with food deficit, 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 
and the martens took them off (V.A. Popov). 

In the southern parts of the species range, marten in the snow- 
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 snow- 
free 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 other food is found in 
great quantities. There, when nut pine is absent in the taiga, ash 



860 

fruits and other plant food appear in marten food in winter when 
principal foods are 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 Cau- 
casus, together with a whole list of wild fruits (including moun- 
tain) ash, those of yew, which are poisonous to man, are of essential 
importance. 

Polyphagy is a very characteristic feature of the pine marten. 
It enables it, in case of a deficit of one food, to turn to another. 
However, the comparison of combinations in one stomach of vari- 
ous numbers of foods with the degree of fullness of the stomach 
and the fatness of the animal, shows that differences in daily ra- 
tions is a negative feature, indicating insufficiency in the main 
full-value foods. The analysis of multiyear data on food of pine 
marten also shows that along with the existing relationship be- 
tween 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 demand for another food 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- 
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 avail- 
able food assortment. At the same time, the occurrence of the more 
difficult to obtain foods falls off: squirrels and tetraonid birds. It 
has been already mentioned above that apart from this general 
situation, there are some local deviations. Thus, in Arkhangel'sk 
district (Grakov, 1964) in summer food voles were found to 
constitute 70% of the data, birds— 23.2%, insects— 24.2%, fruits 
577 (including berries) — 21.2%, and among birds, the importance of 
tetraonids declines and the role of the small birds grows, lizards 
also appear, and the role of shrews — a food of last resort and 
squirrel decreases. 



861 

The stomach of marten holds a quantity of food equal to 1/10 
of the live weight of the animal — this is the optimal daily norm in 
nature. It is not often achieved. The fullest stomach contained 60- 
90 gm of food, but more often 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 stom- 
ach 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 aver- 
age 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 encoun- 
tered. 

Home range. From the time of dispersal of the young, the pine 
marten spends its whole life 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 litter usually remain 
until 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^ 
to 9.2 km^) (Konchits, 1937; Yurgenson, 1937, 1939). In Babkovsk 
forest tract (on the border of Penzensk and Tambovsk districts) — 
4.4 km^ (Yurgenson, 1939), in Zhiguly (P.B. Yurgenson), 4.0 km^ 
in the montane forests of the northwestern Caucasus, the average 
area of the daily home range (11 cases) is 46.7 ha (Donaurov, 
1949; Ryabov, 1959), in fir forests — from 32 to 72 ha. In certain 
cases, martens lived in isolated forest islands with an area of from 
90 to 300 ha. 

The territory of the home range is assimilated by martens in 
succession, and the same range may itself comprise places periodi- 
cally visited by the marten in different hours of its diel activity 
period, or else, as its hunting area. Individual parts of the entire 
home range are not used for hunting and only serve for movement. 

Burrows and shelters. The most preferred shelter of the pine 
marten is 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 
of 30 cm and more. The opening of the inhabited hole mostly 
occurs high above the ground (from 2-2.5 m — 4-5 m; in the north- 
ern Caucasus — at a height of 10-20 m and more), but in certain 
578 cases, the entrance to the hole is situated in the main trunk; 
the presence of light-penetrating cracks in the hollow is 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 conifer- 
ous 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 in thick snow where it is safer and warmer. In Arkhangelsk 
oblast (Garkov, 1964), also in the first half of winter, they more 
often (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 
shelters, 11 were located in hollow fir trees, 2 were in wind-felled 
trees, and the rest were in hollows of beech, aspens and in wind- 
falls. Nests of litters in this region always occur in tree hollows. 
In holes with litters bedding, except the remains of rotten wood, 
does not occur. In the Tatar Republic, the marten sometimes occu- 
pies logs placed in the forest for bees especially in second-growth, 
where hollows are few (B.S. Popov). Very rarely, pine marten 
establish themselves in rock clefts (Zhiguli). 

In the winter time, small stores of food are sometimes discov- 
ered in holes. In such a way, holes inhabited by bees may be 
considered unique *'larders" of this animal; the marten feeds on 
their honey for a long time, making beaten tracks to such holes. 

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 
young pine marten, however, can sometimes be seen in the day- 
time. 

Leading a semi-arboreal way of life, the pine marten climbs 
trees, but not always and not everywhere. Its occurrence and move- 
ment 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 
feed mainly on voles, only 1% of its diel movements were among 
trees (of 132 km). In the second half of the winter, this occurs 
somewhat more often (Grakov, 1964). There, where food of the 
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 
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. 
579 The character of pine marten movement in winter is deter- 
mined 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 diel movements (tracks) is determined by the 
abundance and 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. 

As with area of the home range, the length of diel movements 
(tracks) are closely connected with the quality of land, with its 
food supply, and regularly decreases from north to south. In the 
forests of the Kola Peninsula, it ranges from 10 to 25 km (16 km 
on average) (Nasimovich, 1948); in Priozersk region of 
Arkhangel'sk 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 food- 
rich 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 
forests of the southwestern part of Kalinin district — from 0.6 to 
5.5 km (3.1 km on average; Yurgenson and Konchits, 1937), and 
finally in the Caucasus preserve — from 0.6 to 3.2 km (2 km on 
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 
580 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 



866 

to move in search of suitable, unoccupied areas. In particular, 
young relocate to habitats temporarily rich in squirrels 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 dispersion of young, expansion of 
the range may proceed. 

Reproduction. Up to now, the time when pine martens attain 
sexual maturity is 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, on July 18, the testes were found to be greatly 
enlarged. About 45% of all females in a population bring forth 
litters. In the Caucasus, the percentage of barren females in the 
third year of life and older is 18.7%. For Arkhangel'sk district, 
barren females comprise 22% (Grakov, 1964). 

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 is delayed until the end of 
August. It commences somewhat later than in sable. 

Generally, pregnancy (Manteifel', 1934; Starkov, 1947) lasts 
236-237 days; from other data (11 cases), 254-256 and 258-274 
days (Starkov, 1947). In kidas*, it continues 280 days (Portnova, 
1941). Pregnancy extends through a long period of inhibited 
development (latent period). Its duration is variable. As a rule 
placetation, being connected with the resumption of sexual activ- 
ity, and which is suppressed by September, takes place in March. 
The so-called false rut is dated to that time, i.e. an increase in 
excitement of the sexually mature animals which at that 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 mm in length were found in 
martens killed on 9 and 20 January. This situation, and the pres- 
ence of sexually immature individuals, explain why the false rut 
does not occur in all martens. A marten impregnated on 15 July 
in the German Federal Republic gave birth on 5 January after 
being transported to Argentina; the duration of pregnancy due to 
the latent period was shortened to 176 days (Ulrich, 1953). 

*See pg. 833. 



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 in April (Starkov, 1947), or 
22-26 March (Manteifel', 1934). In case of early placentation, 
parturition should take place in February. 
581 The number of young in a litter ranges from 2 to 8, most often 

3-5. Average size (for entire range) is 3.8; in the Pechora basin, 
it equals 3 (2-4), in Moscow district, 5 (3-7). In the northwestern 
Caucasus, litter size ranges 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). 

Growth, development, and molt. Growth and development of 
the pine marten take place as in the sable. 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, fe- 
males begin to fall behind males in weight: at the age of 10 weeks, 
animals weigh 400 gm, at 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- 
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 devel- 
ops. 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. Dan- 
gerous 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 
the 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 
infection and the epizootic caused by it were noted in Severnii 
Territory in 1951/52-1955/56 when the number of martens was 
high. 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 1953- 
1954. 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 more heavily than adult males and the 
yearling of both sexes. Poorly nourished martens are always more 
strongly infected. 

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 may be 
considered as competitors of pine martens — red fox, forest wildcat 
[Felis silvestris], sable, stone 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 Pechora- 
Ilych 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 
three years — 2.7%. In the same preserve, from material of com- 
mercial hunting for 1937/38-1948/49 (309 specimens), yearlings 
constituted 49.9% on the average (from 44% to 58%). In the fol- 
lowing 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, in the time of the harvest, a 
larger percentage of yearlings is usually captured than their actual 
proportion in the population. Of 464 martens commercially taken 
in the 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 mar- 
tens, the sex ratio was 1.0 : 0.4 and among those up to one year it 
was 1.0 : 1.7. On the whole, males in Pechora population consti- 
tuted 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 (1 
year and older) draws attention to itself. 

Of 126 martens taken in 1939/40 in Volzhsko-Kamsk Territory 
(Aspisov, 1959), juveniles constituted 51.6%, in their 2nd year 
[yearlings] — 30.2%, in their 3rd year — 11.9%, in their 4th year — 
3.9%, and in their 5th year — 2.4%. Individuals above five years 
old were absent. In 1940/41, among 289 harvested marten, were 
juveniles — 70.6%, yearlings — 19.4%, in their third year — 6.2%, in 
their fourth year — 2.4%, and in their fifth year — 1.4%. It is quite 
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 in this 
commercial catch always and everywhere predominated. 



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, 
the sex ratio in the population changed in favor of females (from 
38% to 56%). In the unfavorable years, the percentage of pregnant 
females fell as much as 83% (Kirov district, 1959/60). This was 
also observed in 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 per- 
centage of juveniles is 48% with annual fluctuations from 35% to 
58% (Teplov, 1954). 

In Arkhangel'sk district, during 13 harvest seasons (1949- 
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- 
lished (Grakov, 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 
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- 
tuted 55.2% (ranging from 49.9-58.2%), yearlings— 31.7% (29.1- 
38.6%) 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 fe- 
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 
several successive seasons combined. The rarity of such combined 
events determine the rarity of noticeable population changes. 
Usually, deficiency of one main food 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 north- 
ern Europe was observed in years of favorable feeding conditions, 
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- 
est number of dead animals was found in 1950/51 and 1954/55. 
The next rise in numbers was in 1957/58 (Gribov, 1959; Grakov, 
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. With intensive harvest- 
ing, 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, i.e. for 10 years, the frequency of occur- 
rence of their tracks in pine forests along a 10-km route in winter 
fluctuated between an average of 0.5 to 0.95; i.e., by 1.9* fold and 
in the region of dark coniferous forests, more favorable for the 
pine marten — from 0.8 to 1.3; i.e., only 1.6 fold. In Arkhangel'sk 
district (Grakov, 1962) during eight years (1952-1959) this fre- 
quency fluctuated from 0.7 to 2.9; i.e., 4.1 fold. 

It is 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 district, the pine marten population is esti- 
mated to be plentiful. 

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 litters. 
584 Field characteristics. It is rare to encounter a pine marten. 

One may often observe the characteristic feces of pine marten on 
logs, stumps or on forest paths — sausage-like in form, length 8-10 
cm with diameter about 1-1.5 cm. Usually, it is 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, the tracks of pine marten are not clearly printed due 
to the dense fur on the food pads; in the stone marten, the callosi- 
ties of the feet are always obvious in the footprints, even in winter. 

The most typical gait of the pine marten is a bound, leaving on 
the snow an extended chain of paired footprints. 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 
is easily determined 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 jump- 
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 exceeded 
only by that of sea otter, beaver, and sable. Its fur is durable, 
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). 

Exhaustion of their stock as a result of overhunting has led to 
the necessary prohibition of its hunting everywhere. This measure 
was shown to 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 are: 
1) 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 
of a dog, is the most effective one, but for this, good dogs are 
essential. The period of harvest must be restricted to two months: 
December-January. The effectiveness of the various methods of 
capture for pine marten is only known for the northwestern 
Caucacus (Ryabov, 1958). Tracking after the first snowfall is most 
effective, especially in deciduous forests. Hunting with the aid of 
a laika dog comes in 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 them- 
selves widely. 

With the regulation of harvest, the number of pine marten is 
easily maintained at a high 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.). 

STONE MARTEN38 

Martes (Martes) foina Erxleben,** 1777 

1911 . Mustela foina. Erxleben, Syst. Regni Anim., 1, p. 458. 
Germany. 



♦Special sorts of traps — Sci. Ed. 

^^Also, belodushka (white-breast), " kunitsa-belodushka" (white-breasted marten) 
and among furriers, kunitsa gorskaya (mountain marten). 

♦♦Misprinted in small lower-case type in Russian original — Sci. Ed. 



875 

1192. Mustela domestica. Pinel. Actes Soc. H.N. Paris, 1, p. 55. 

France. 
\?>Q\. Mustela foina alba. Bechstein. Gemeinn. Naturg. 

Deutschlands, 1, p. 759. Thuringia, Germany. 
1869. Mustela martes var. fagorum. Fatio. Faune vertebr. Suisse, 

1, p. 318. Switzerland. 
1873. Mustela intermedia. Severtzov. Vertik. gorizont. raspredel. 

Turkestansk. zhivotnykh. Izv. Obshch. lyubit. est., antrop. 

etnogr., 8, 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, 2, p. 

120, tab. 2. Tbilisi. 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. 1, p. 111. Altai. 
1917. Martes rosanovi. V. et E. Martino. Zap. Krymsk. obshch. 

estestvoisp. lyubit. prirody, 7, 1 (reprint). Northwestern slope 

of Chatyr-Dag, Crimea (Crimean preserve; V.H.). 
1946. Martes foina ognevi. Laptev. Izv. Turkm. fil. AN SSSR, 2, 

p. 57. Central Kopet-Dag, Bol'shoi Balkhan (type and type 

locality not indicated). It is 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 length. Color 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 
relatively short and widely separated. Longitudinal diameter of the 
inner half of the upper molar is only a little larger than the outer 
half (V.H.). 



•■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 many 
individual features, to the pine marten. It 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 
their inner edges is about 47 mm, in the stone marten — 54-56 mm; 
Schmidt, 1943). In stone marten, the bare tip of the nose is light, 
usually of light flesh-color or grayish; in the pine marten, it is 
dark-black or grayish-black (Schmidt, 1943). All these character- 
istics 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 is more noticeable. The stone marten stands and moves 
in a manner which differs considerably from that of the pine mar- 
ten — they often appear to be "creeping" like the polecat, and do 
not bound ("gallop") as do the pine marten and sable. This, evi- 
dently, depends also on the fact that the length of the forelegs 
(height at the "withers") differs in each species — in the pine mar- 
ten, it is 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 
marten, but is somewhat lighter. The underfur is also lighter — not 
grayish, but whitish. The tail is dark-brown; the color of the back 

587 is darker than that of the pine marten. In the remainder, its color 
is like that of latter. 

The differences in color and form of the throat patch of both 
species are very sharp. In the stone marten, it is always white, and 
only in the form of a rare exception has it light pale highlights. It 
is large and, as a rule, two projections extend backwards to the 
base of the forelegs and also extends upward on the legs, some- 
times halfway. Thus, the dark color of the belly juts out between 
the forelegs as a line into the white color of the chest and some- 

588 times 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 unnotice- 
able 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 University (upper row, after Wehrli, 1932*, with modifications). 

*Not 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 localiza- 
tion is observed. 

There is no sexual differences in color and character of the 
fur. Young animals in their first autumn have fur not differing 
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 sub- 
orbital 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 — a feature better developed in adults and older individuals). 
3. Nasal bones have a well-marked constriction ("isthmus") in their 
middle part (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 
form an angle). 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 
their middle part). 6. Projection at anterior edge of sphenopalatine 
notch absent or weakly defined. 7. Upper carnassial tooth some- 
what larger (its length usually more than diameter of upper molar 
lying next to it). 8. General dimensions of upper molar relatively 
smaller and its inner blade considerably smaller than in pine 
marten. In linear measurements, this blade is only a bit larger than 
the outer one, rarely equal or almost equal to it. Its longitudinal 
diameter is considerably less than diameter of entire tooth. 9. On 
outer (lateral or posterolateral) surface of upper molar, there is a 
clearly defined vertical fissure (this tooth surface is evenly convex 
in pine marten). 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, 6, 
pelvic, 3. Clavicles are smaller than in pine marten; their length is 
about 10 mm. As in pine marten, they are flattened, but they are 
all 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. The penis of the stone marten is 
590 larger — its preputial part is about 33 mm (Shtreili, 1932). The 
form of the os penis is the same, but larger and massive in its 
dimensions (see below); the os penis of a young stone marten is 
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 well- 
marked 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), dimen- 
sions 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. 

Condylobasal length of skull of males 71.0-86.0; of females 
73.2-82.8 mm; zygomatic width of males 42.2-56.4, of females, 
43.6 to 50.6 mm; height of skull of males 30.1-32.6, of females, 
29.1 to 31.0 mm (measurements of skull after data of Kuznetsov, 
1941, 31 males, 29 females). 

Weight of males in winter is 1700-1800, in summer the aver- 
age is 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 1325- 
1950 gm, of females 865-M 1060 gm-1306 gm (20 specimens in 

*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 
long time 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 50.6- 
M 54.8-58.2 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 
males. In body length, this difference is not, however, more than 
100 mm (in the Central European martens). Dimensions are sub- 
ject to negligible geographic variations. On the whole, variation in 
meristic characters within the USSR, as in color, is less than that 
in the pine marten (V.H.). 

Systematic Position 

The stone marten is a well-defined species. Regardless of signifi- 
cant similarity of features, it 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 [yellow- 
throat marten] (M. flavigula) and il'ka [fisher] (M. pennanti), are 
clearly divided into two groups — the stone marten constitutes one 
and all the remainder; i.e. sable, pine and American martens (M. 
americana), even, if they are considered separate species — the other. 
It 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 infor- 
mation 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, combin- 
ing 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 combina- 
tions are encountered. In such cases, accurate identification is only 
possible from the skull. 



883 

Only one, not fully defined, case is known, of a hybrid off- 
591 spring, which 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 male sex organs (see above in corresponding places 
in "Description" sections of both species). 

The stone marten is a more ancient species compared to the 
pine marten. It is, apparently, a Pliocene form. Both species are 
strongly differentiated from each other by their habits and ethol- 
ogy (Schmidt, 1943) (V.H.). 

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 
of this part of the range are not well-understood and reliable data 
on them are scarce. In the north, the boundary starts at the Gulf of 
Finland west of Leningrad, apparently somewhere on the meridian 
of Chudsk Lake (the marten is known from Rakvera, the former 
Vezenberg), and extend southward from western Estonia, and 
Lithuania. 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 — Surazha, Pochepa, Trubchevska, and Pogara regions (new 
data of A.V. Fedesov — but according to Fedosov and Nikitin, 1951, 
this species is not recorded in Bryansk district; Melander, 1938). 
It is not excluded that this marten lives in the southern parts of 
Mogilevsk district. 

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, 
the boundary passes to Bobrov region southeast of Voronezh 
(Sadovoe; Ognev and Vorob'ev, 1924). South of Voronezh, the 
stone marten was recorded in the upper Tikhaya-Sosna and 
Chernaya Kalitva (Budenovsk and Ladomirovsk regions; Barabash- 
Nikiforov, 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 de- 
scends 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 pre- 
sented 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 its 
"established" occurrence in Moscow district 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 
Chudsk Lake to the upper Volga, and along it [river] to Zhigulei 
and nearly to Saratov (map), and then to Dnepropetrovsk and along 
the Dnepr to the Black Sea. 
593 In the literature there are a series of precisely known 

occurrences of stone marten outside the above-outlined line. This 
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 
governances, if these areas are considered as corresponding to the 
contemporary districts, are interpreted as follows. The most west- 
erly 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 
sporadic inhabitation are possible. The above described border of 
the range actually passes along the border of this district. The 
existence of the described species in Smolensk district still is not 
established. 

From all that has preceded, particular attention should be given 
to information on occurrence of the stone marten in Moscow dis- 
trict, along its southern outskirts and at its border. Old and poorly 
defined records (Sabaneev, 1872) for the district at that 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 I have not succeeded in learning anything 
positive about the existence of the marten, M. foina Briss. with us (middle Volga area; 
V.H.). It 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 it is not possible 
to consider them confirmed. However, neither can it be fully ex- 
cluded that the stone marten may rarely and sporadically be found 
somewhere in the Tula district (it 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 per- 
manently somewhere here. Captures of stone marten in Ozeri is 
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 from the periph- 
eral part of the range absolutely should also be based on skull. 
Insufficiently critical consideration of reports on the genus led to 
misrepresentations in revisions (Ognev, 1931; Novikov, 1956 and 
particularly Yurgenson, 1933). 

Some corrections to the boundary of the range of marten are 
possible, but at the present time, only that outlined above is reliable. 

In the Crimea, the marten is widely distributed in the moun- 
tainous part of the peninsula, and is apparently not found in the 
plains at a distance 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; i.e. the shore of the Caspian Sea. Along the valley 
of the Terek, the marten extends east of Grozny. Along the Cas- 
pian coast, it reaches or almost reaches the sea through the gallery 
forests of the Samur [river] as far as its mouth; it is encountered 
on the Apsheron peninsula (Heptner and Formozov, 1941; 
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, it was met with there 
(Satunin, 1915). 



888 

594 The stone marten is present in Talysh. This area of occurrence 
within the USSR is, possibly, isolated from the rest of the Cauca- 
sian range. 

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 
Bol'shoi Balkhan [range] (Laptev, 1934; V.G. Heptner). In south- 
ern Turkmenia, it is encountered in Gyaz'-Gyadyk (right 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. 

The section of the range lying beyond the Amu-Dar'ya is quite 
large and includes all the mountain ranges from the Pamiro-Alaisk 
to the Tien Shan systems. Westward the range extends apparently 
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]. It also inhab- 
its 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 separate part of the range occupies southern Altai. The mar- 
ten here is distributed southward to the Kurchum, Bukhtarma and 
Ul'ba [rivers] (Sludskii, 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 the Chulyshman (Yurgenson, 
1938; V.G. Heptner). In the Sayan and in Tuva, particularly in 
Tanna-Ola, the marten is 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 a 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 oblast. 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 in- 
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 Ire- 
land) 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 Tibet,'^° probably Shansi and Chihli (Hebei), i.e. 
the region between the Huang He, the ocean and the Great Wall. 
This does not exclude the possibility that it 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 north- 
western 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. 

In the literature 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 
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 re- 
liable 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 in Tibet are known. It is very probable that it does 
not occupy the whole country. There is information for Kam and Nan-Shan (Ognev, 
1931). The basis for confirmation of its occurrence in the Nan-Shan is not clear. 
Reported for Lhasa. 



891 

also in the Great Khingan according to Lukashkin and Zhernkov, 
1934). The reports of Sowerby (1923), whose mistakes are 
recognized, are not reliable'*' (see also range description of pine 
marten, page 846), or may be based on individual variants of Man- 
churian sable, the quality of which is 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 
presence here (Shansi and Chihli-Hebei) calls for skepticism*. In 
any case, it is now generally believed that in northeastern China, 
at least in its northern (Great Khingan, Il'khuri-Alin') and eastern 
parts (eastern Manchurian mountains) martens are absent. The whole 
question of the existence of stone marten in China demands spe- 
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). 

In the zoogeographical literature, the stone marten is usually 
evaluated as "European", or "western" form. As seen from the 
outlined range, it is typically and Near- and Central Asiatic, and in 
part a "Western Mediterranean" species closely associated with 
mountains. Only, secondarily, apparently, did it settle in Europe. 
Its occurrence in the European plain far from the mountains, but 
not in other places, is apparently connected to a significant extent 
with humans and their constructions. In Central Europe, it is a 
well-marked synanthropic species (V.H.). 

Geographic Variation 

Geographic variation of the stone marten within the USSR is in- 
significant. This is related to the amplitude of racial differences, as 
well as the number of forms. It is highly probable, that their number 

■"This author, for example, states that "It is abundant in the western portion of 
the country (former Manchuria; V.H.), whence its 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. 
68)**. 

*Presence confirmed by Zhang et al., 1997 — Sci. Ed. 

**In the copy I have consulted, this quote is on p. 66, paragraph 3 — Sci. Ed. 



892 

is less than what is usually considered. The fur industry differen- 
tiates 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 be- 
long. Evidently, geographic variation in that part of the range which 
lies 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 (syn. 
domestica, alba, fagorum). 

General dimensions quite small, skull of average size. 

In winter fur, color of back varies from quite light grayish 
tawny to completely dark brown. Guard hairs are tawny or chest- 
nut-brown, underfur is very light, pale-gray or whitish, sides a bit 
lighter than 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 in size and 
form. 

Body length of adult males usually 450-500 mm; of females, 
400-440 mm. Tail length of males— 250-270 mm, of females 230- 
250 mm. 

Condylobasal length of male skull (9) 71.0-M 82.00 ± 0.65- 
84.6 mm; 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 47.8- 
50.0 mm; height of male skull 31.0-M 31.9 ± 0.49-32.6 mm. 

Present in European part of the USSR, except Crimea. 

Outside the USSR — in western Europe, except the Balkan 
Peninsula and Pyrenees [Iberian] Peninsula. 

The marten of the part of our country mentioned belongs to 
the nominal form, so far as is 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 
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. et E. Martino, 
1917. 

Dimensions somewhat smaller than in preceding form. Skull 
small, not massive. 
597 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 size. 

Body length of males 465-510 mm, of females (1) 470 mm; 
tail length of males 300-325 mm, of females — 275 mm, length of 
hind foot of males 94-95 mm; height of ear of males 43-47 mm, 
of females — 41 mm. 

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 43.8- 
48.1 mm, of females 42.6-M 45.7-47.0 mm. 

In montane Crimea. 

Outside the USSR, absent. 

The Crimean form is very near to that described above, and is, 
apparently, identical with it (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 ten- 
dency 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 

■•^Measurements of skull, as in other cases, given after Kuznetsov (1941), figures in 
parentheses after M — 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)- 
56.4 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 
females 29.1-M 29.7 ± 0.14 mm (28.9 mm— 30.4 mm). 

For dimensions of os penis, see above, "Description" section. 

In Caucasus. 

Outside the USSR, probably present in contiguous parts of 
Iran and Turkey. 

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 49.9- 
M 51.4-52.0 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 
to Tarbagatai and Altai. 

Outside the USSR, in northern Iran (?), 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) differ also from the eastern (Tien Shan); however, the dis- 
tinction of the form ognevi was presented without adequate 
argument. Identity of Himalayan and Tien Shan martens is estab- 
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] Penin- 
sula; 2) M. (M.)/ bosniaca Brass, 1911 — Yugoslavia; 3) M. (M.) 
/ milleri Festa, 1914 — Island of Rhodes; 4) M. (M.) f. bunites 
Bate, 1906— Crete; 5) M. (M.)f. syriaca Nehring, 1902— Syria; 6) 
M. (M.) / toufoeus Hodgson, 1842— Tibet, Lhasa; 7) M. (M.) / 
kozlovi Ognev, 1931 — Kam, eastern Tibet. The last two forms ap- 
parently are synonyms and possibly belong to intermedia (V.H.)- 

Biology 

Population. Information is scanty. The greatest abundance of stone 
marten is in the Caucasus and Trans-Caucasus. In Middle Asia, 
they are encountered more rarely, because habitats typical for them 
are not found everywhere and are unevenly distributed. It is not 
rare in the 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 
is 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 forest- 
steppe, Novikov, 1962). In the middle [forest] zone, it is a great 
rarity. 

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 sig- 
nificant extent into populated areas, in the role of a synanthropic 
species. Here, it prefers to live in stone structures — in attics, cel- 
lars, sheds, stables, schools, storehouses, railway stations, in stone 
walls, mill weirs, and even in old stork nests, etc. 

In the central forest-steppe (Belgorod district), the stone 
marten (Novikov, 1962) is a characteristic inhabitant of forests. It 
prefers old tall-trunked oak-groves on hills, especially those with 
broken relief. It more rarely settles in the middle-aged mixed for- 
ests or oak-groves, and avoid the sections with low relief. Here, it 
also settles in buildings. 

Primary habitats are characteristic for those parts of the range 
which exhibit a more or less distinctly montane character. They 
are distinguished by less harsh climate; snow cover here is not 



896 

permanent and is uneven. These are unforested or almost unforested 
mountains, ravines, rock slides and breccias with, characteristi- 
599 cally, shrubby vegetation of a Mediterranean type, "maquis" or 
sheblyak*', forests and rocky and stony mountain slopes, deserted 
orchards, vineyards, and nut groves. It does not avoid montane 
forests, but in the Caucasus, it clearly prefers broadleaf forests in 
foothills. It avoids the dark coniferous forest zone, mainly because 
of their snowy winters. It is also encountered in rock outcrops and 
slides at the upper forest limits and even above, up to 2600-2700 
m 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 1 cm^ of the supporting surface of the stone 
marten foot is double that of the pine marten (Ryabov, 1959): 30.9 gm 
and 15.2 gm; (Ryabov, this is why it obliged to avoid snowy regions). 

Food. A typical feature, distinguishing the stone marten from 
pine marten and sable, is the 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): 



Vertebrates 


76.0 


Plant foods 


62.3 


Mammals 


59.0 


Olive fruit 


21.6 


Rodents 


55.6 


Blackthorn fruit 


19.9 


Birds 


44.6 


Wild pear fruit 


8.5 



♦Russian word for shrubby deciduous successional vegetation — Sci. Ed. 



897 



Sparrows 


34.1 


Sunflower seed 


11.8 


Lizards 


7.0 


Common cherry fruit 


3.5 


Insects 


50.0 


Malagensk cherry fruit 


4.5 


Beetles 


50.0 


Mulberry fruit 


4.1 


Insect larvae 


4.4 


Rose fruit 


7.4 



Among vertebrate animals, the following were found: 

Common wood mouse 16.2 Hares (young) 13.8 

Gray hamster 14.2 Mole rat 6.0 

Common vole 7.0 Spotted ground squirrel 5.4 

Kurgan [barrow] mouse 1.4 Common hamster 2.2 

In summer, rodent occurrence in the food of martens reached 
75-88%, and insect occurrences — 75-81%. Food composition 
changed significantly annually, depending upon fluctuations in 
natural abundance of various foods. 

In the "Vorskl Forest" (463 specimens; majority pertain to 
autumn and winter) (Novikov, 1962) rodent occurrence in various 
years and seasons varied from to 75%; in summer from 20 to 
75%, in autumn from 1 1 to 33%, in winter from to 57.9%. The 
red vole was most often encountered: for the whole period, 7.3%, 
in summer, 16.1%; the subterranean vole — 0.9%, and in summer 
12.8%. Also found were water vole, the yellow-necked field mouse 
and house mouse, and field hare (in winter up to 1.5%). 
600 The occurrence of birds in the marten diet varied in summer 
from to 33%, in autumn — from to 16.7%, in winter — from 
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, 
1962). However, individual animals sometimes systematically at- 
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 
to 100%, in winter — from to 33.3%. Among insects, beetles 
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 
winter. In summer, the fruits and berries occurred from to 70%, 
in autumn — from 9.17 to 100%, and in winter — from to 100% of 
the cases. Occurrence of sloe was 36%, pear — 42.7%, and apple — 
17.0%. No less than 27 species of plant food were revealed. The 
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 
marten feeds both on carrion, as well as fruits hanging from the 
branches of trees and bushes. 

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 
found in 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 
Shikina, 1936). 

In the Zakatalo-Nukhinsk valley (Azerbaidzhan), the food of 
the stone marten was composed of the following elements 
(Rukovskii, 1957; % of occurrence). 

Mammals 

Birds 

Reptiles 

Insects 

Acorns 

Blackberry 

The proportion by weight of the insects and plant food here is 
also quite significant. 

In montane Crimea, the stone marten eats grapes and grape 
snails (Flervov, 1929). According to analysis of 373 specimens in 
the Crimean preserve in 1936-1938 (M.I. Savvina) the food com- 
position of the stone marten was as follows: 

Mouse-like rodents 41.0 Molluscs 0.9 

Hares 1.3 Pears 19.0 



65.5 


Grapes 


10.8 


5.1 


Dogwood 


16.1 


3.5 


Myrobalan plum 


3.5 


84.0 


Blackthorn 


4.3 


2.4 


Pear 


3.5 


23.3 


Nuts 


1.0 



899 



Roe deer (carrion?) 


1.0 


Dogwood 


7.1 


Birds 


16.0 


Rose 


9.3 


Lizards 


0.3 


Mountain ash 


0.3 


Fish 


0.5 


Juniper 


0.5 


Insects 


50.4 


Mushrooms 


1.8 


Insect larvae 


9.1 


Conifer needles 


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 Europe (Brehm, 
1866; Chudi, 1873) refer to rats, 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. 

Young martens in captivity ate in 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 yellow- 
necked mice, weighing 20 gm each. Thus, the daily ration was 
about 170-250 gm of animal food (Abelentsev, 1958). In summer, 
a young marten ate in one day about 15 apricots whose flesh 
weighed about 260 gm. Upon receiving animal food, stone martens 
did not refuse plant food. In hot weather, they ate only mulberry 
fruits, cherries, etc. In captivity, plant foods were often preferred; 
when offered each day 10 mice and 250 g of apricot flesh for 
several successive days, they ate the fruits first and left the mice. 

The ration for martens in the nursery [animal facility] of the 
Zoological Institute of the Academy of Sciences of the USSR 
consisted of 150-180 gm of rabbits, 25-30 gm of milk, 20-30 gm 
cornelian cherry or briar fruits or 30 gm boiled carrots per head. 
On such a ration, the martens felt quite satisfied. 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 deter- 
mined 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 
(Abelentsev, 1958), the maximum length of the marten's daily track 
was 5 km, but sometimes a nursing female did not go far from her 



900 

den. Tracing of 16 daily tracks in "Vorskl Forest" in Belgorodsk 
district showed that the average length was equal to about 1 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 1100 m. The daily 
track may sometimes be in the form of a closed loop, but, more 
often, it extends in any direction. In one case, the daily range of 
activity 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 unin- 
habited 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 
(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 is inferior to the pine mar- 
ten in its ability to climb trees. However, it climbs well and in 
dense forests (Crimean preserve; V.G. Heptner), it does so fre- 
quently. It is a capable swimmer. It is active by day most often in 
602 summer (Dinnik, 1914), when the nights are short. In the central 
forest-steppe (Belgorodsk district), the stone marten leads a 
crepuscular-nocturnal way of life, but in summer it is observed by 
day, even in midday hours. Martens living in open-air enclosures 
were outside at 1700-2000 hours but hid in the nest in the early 
morning— by 0600-0700 hours (Novikov, 1962). 

*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 sur- 
face. Result from 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 
heaps, overhanging turfs, holes, root-hollows, etc. Martens also 
dug in the snow 34 times in open places, and near trees 23 times. 
For a considerable part of way stone marten went on foot, not 
lingering, and making only small turns. Only in thickets of shrubs, 
their tracks are found to meander, a characteristic for the period of 
fattening. 

In soft snow, martens prefer to use for their movements paths 
of hares and ski-tracks. They rarely run along their own tracks. 
Martens usually move in large jumps measuring 30-40 cm in length. 
On firm or crusted snow or, the gait 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 in the 
grass, they bound (Novikov, 1962). 

Seasonal migrations and transgressions. Information is absent. 

Reproduction. Estrus and copulation, according to the observa- 
tions of fur breeders, occur at the 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 
603 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 
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, 
1962) the young 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 popu- 
lation dynamics. The pine marten, and perhaps the red fox may be 
considered enemies of the stone marten. Large diurnal and noctur- 
nal birds of prey, wild cats and other carnivores are a danger to the 
young stone marten. A case was described when a pine marten 
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 pos- 
sibly 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 
from the stone marten (Greve, 1909). 

Longevity of the stone marten is not established, but is, prob- 
ably, similar to that of the pine marten. 

Concerning population dynamics, there are almost no data. 
Among the three species of the genus of true martens, genus Martes, 
it is the stone marten which most easily adapts to changes intro- 

604 duced 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. When encountered in nature, usually very 
briefly, it is difficult to sort out those features which distinguish 
the stone marten from the pine, the more so since, in some places, 
they are met with in similar habitats. Non-forested places or those 
in immediate proximity to human habitation is the best indicator 
that it is a stone marten. It is also said that finding the animal or 
its tracks among rocks and talus is 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 is a quite valuable fur animal, but is inferior, not 
only to sable, but also to pine marten. Its share in the fur market 
of the USSR is not great: stone marten constitute no more than 10- 
12% of that of processed pine marten. Its 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 
villages there, living in bell towers and other buildings (Manteifel', 
1947). 

The stone marten is captured with jaw traps, box type traps for 
live capture, and various sorts of [kulemka] and plate [plashka] 
traps. 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 some- 
times 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. p. 88. Nepal. 
\^\\. Mustela aterrima. Pallas. Zoographia Roso-asiatica, 1, p. 

81. Between the Ud and Amur rivers. 
1862. Mustela (Martes) flavigula var. borealis. Radde. Reisen Slid. 

Ost. Sibirien, 1, 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 a true marten. Its typical 
martenness is 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 
2/3 the body length. The tail is covered with short hairs, much 
thinner than in all other martens, and therefore seems even longer 
than it actually is. The limbs are strong and relatively short with 

■•'The proposed change of this name to Lamprogale Ognev, 1928 (see page 749) 
did not find wide application among systematists because it was not called forth from 
actual requirements and did not correspond with nomenclatorial rules. 

■•^The kharza is 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. 



906 



broad feet — in moving, the animal strongly bends its back into 
a hump and moves in bounds which, at a rapid pace, are very 
long. 

The kharza in 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 — even more 
so than the stone marten. The ears are large — broad, but short, and 
are 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, in general, the paws are weakly furred. In 
short summer fur, the kharza appears still longer and thinner, with 
a long, thin tail. 

Winter pelage differs sharply from that of our other martens in 

that it is relatively short, and, moreover, is 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. Young female kharza Martes (Charronia) flavigula Bodd. "Kedrovaya" Pad 
preserve, southern Primorye. 18 December 1964. Photograph by A.G. Pankrat'ev. 



907 



species of the genus. The top and occiput of the head is blackish- 
brown 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 is the hair 
on the ear margin. Starting at the occiput along the dorsal surface 
of the body, the fur is a shiny brownish-yellow color with a golden 
tone. Posteriorly, this color gradually becomes browner and the 
rear portion of the back, and the region of 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 
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 tail hair is grayish-brown, the 
claws are white. 

Individual variation in the fur 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 summer fur is somewhat duller and 
darker — the golden tones along the back are more weakly devel- 
oped. The fur of young martens is somewhat lighter than that of 
adults. The black tones are less pure, with a brownish mixture and the 
throat patch is not so bright. There is no sexual differences in color. 

The skull of the kharza in general appearance, and in a series 
of details, is more similar to the skull of the stone marten, differ- 
ing from it, most notably, in its large dimensions (see below). The 
brain case is relatively shorter — width between the mastoid 
processes is equal to, or is a little greater than the distance from 
the lower edge of the occipital foramen to the posterior edge of the 
sphenopalatine notch (ratio is reversed in stone marten). The nasal 
region is broader and shorter. There is no isthmus in the middle 
length of the nasal bones. The [postorbital] constriction of the 
skull behind the supraorbital process is relatively weak and is equal 
to the width of the interorbital space or larger. It is larger, 
sometimes significantly so, than the width of the skull above the 
canines. 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 
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 is less than 
half the transverse diameter of the tooth. The outer lateral surface 
of the upper molar has a vertical groove. 

The female skull is somewhat smaller than the male skull; age 
variation is significant, and generally corresponds to that of 
our other martens. The caudal skeleton is composed of 21-24 ver- 
tebrae (variation apparently, is large). 

The OS penis has a characteristic form and is sharply distin- 
guished from that of our other martens. The basal half of the bone 
is laterally compressed from the sides and is quite tall, and is 
flattened below; a fissure 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 lo- 
cated in the form of a corolla, each at a corner of a quadrant. 
These processes are directed upwards, one of them usually being 
larger than the others (Pocock, 1941 — Indian kharza). 

In all dimensions, the kharza is considerably larger than our 
other martens. Body length of males (9) is 500-M612-719 mm, of 
608 females (12) is 500-M575-620 mm; tail length of males (6) 370- 
M407-442 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) 100- 
Ml 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; 
Bromlei, 1956, with additions). 

Condylobasal length of the male skull (8) 104-M109-112.5 
mm, of females (7) 96. 2-M 100.6-109.4 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 
(7) 37.0-M39.9-46.8 mm (Ognev, 1931; Bromlei, 1956; V.G. 
Heptner). Length of os penis about 76-78 mm. 

Weight of males (eight specimens between October and January) 
is 2463-M33 17-5748 gm, of females (10 specimens between August 
and February) 1155-M2765-3827 gm (Bromlei, 1956) (V.H). 



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 be- 
low). 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 oc- 
cupies 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 53°30' [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 the south'*^ 
Evidence of the occurrence of this species between the Udoi 
and the Amur (Pallas, 1811)* are quite indefinite and may fully 
apply only to those places near the Amur. Information on the dis- 
tribution of kharza in "Amur district north of the south slopes of 
the Yablonovyi range" (probably Stanovyi? V.H.; Ognev, 1931) 
609 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 
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 is improbable. Reference to the 
capture of kharza in the Tuva ASSR ("Uryankhai Territory") on 
the Kemchik [г.] (Ognev, 1931), is completely unintelligeable and 
is obviously mistaken. It is also entirely absent in the Mongolian 
Republic. The source of these mistakes, as in the majority of simi- 
lar 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 (S. Kurg, Nilgiri Hills, Travancore). 
Absent in Ceylon. 

The range of the kharza is very typical of an eastern Asiatic 
Tertiary range. Apparently, the main region for the development of 
this species lies in the south. 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 
described. The majority of them are groundless, and 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 indi- 
viduals are darker). 

The form existing within the boundaries of the USSR appar- 
ently, differs from the nominal (Nepal), although many authors 
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 USSR — in northeastern China (former Manchu- 
ria). The border with respect to the nominal form is not known. 
611 Differences between our kharza and the nominal form are not 

sharp, but apparently are real. Body length of the latter is as fol- 
lows: male (9) average — 571 mm, female (9) — 494 mm; tail length 
of males — 431 mm, females — 408 mm. Weight is somewhat less, 
apparently, and also skull (Pocock, 1941). 



A series forms described in the area lying between the Hima- 
layas and the Amur (kuatunensis, szetchuensis, yuenshanensis, 
melli, koreand) are completely based on information stating its 
similarity to the animal of the Amur form (Jacobi, 1922; G. Allen, 
1938). 

Outside our country, the following forms are usually accepted, 
1) M. (Ch.) f. flavigula L. 1875 — Himalayas, from Kashmir east- 
wards, southern China northward to Shensi and Gansu; 2) M. (Ch.) 
f. gwatkinsi Horsefield, 1851 — southern India (Nilgiri Hills, Kurg, 
Travancore); 3) M. (Ch.) f. chrisospila Swinhoe, 1866 — Taiwan; 
4) M. (Ch.) f. peninsularis Bonhote, 1901 — southern Tenasserim 
and Malacca; 5) M. (Ch.)f. indochinensis Kloss, 1916 — Indochinese 
Peninsula, northern Tenasserim; 6) M. (Ch.) f. saba Chasen et 
Kloss, 1931 — Kalimantan (Borneo). 

Among all 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 
river basin (Yu.A. Salmin and V.D. Shamykin). Northward to 
the extreme limits of its distribution it is rare everywhere. It is 



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 "abun- 
dant". Figures for fur production are not indicative [of population], 
species is not exploited due to difficulty and unprofitability of 
hunting. 

Habitat. Most commonly encountered in montane coniferous 
taiga, although in various habitats — on rocky cliffs with thickets of 
Mongolian oak, in the broad-leaf forest zone, in burnt-over areas, 
in nut pine stands, in montane spruce and spruce-fir taiga up to 
timberline. Most frequently, it lives throughout mossy dark conif- 
erous forests of the Okhotsk type, along the northern slopes of the 
hills — in musk deer habitat — and in the valleys of rivers and streams 
(Bromlei, 1956). In summer, it is 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 
in the arid, unwooded mountains of the Northwestern Frontier region 
of Pakistan (Pocock, 1941). 

Food. Kharza is an omnivorous animal, however, in the Ussuri 
Territory the foundation of its 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 musk deer onto the ice. In 
612 1936 on the Armu and Nantsa rivers, over a distance of 200 km, 
the carcasses of 26 musk deer killed by kharzas were found (1 per 
7.7 km); in 1952, along the Sitsa river — 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). 



916 




Fig. 227. Montane mixed broad-leaf forests in "Kedrovaya Pad' " preserve (southern 

Primor'e) — habitat of kharza, and also Amur badger, spotted deer, leopard and others. 

June 1958. Photograph by A.G. Pankrat'ev. 

Table 61. Seasonal characteristics of diet of kharza (% occurrence) 



Type of food 



Entire year Winter 



Spring 



Summer Autumn 



Musk and roe deer, 

goral, moose calves, 

wapiti, wild pig (suckling) 

Small exploited species 

(squirrel, hare, sable, 

Siberian weasel) 

Mouse-like rodents, pikas 

Hazelhen 

Small birds 

Fish (migratory salmon ) 

Molluscs 

Insects 

Nuts and fruits 



45.5 



27.1 



5.2 
12.6 
3.4 
2.1 
1.4 
0.5 
2.2 



63.1 



29.7 



7.2 



44.9 



26.6 



31.5 42.3 



20.0 32.2 



6.1 


11.6 


3.4 


18.3 


17.6 


6.8 


4.1 


7.7 


1.7 
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 au- 
tumn, fish, fruits and nuts are secondary foods of characteristic 
occurrence. 

In the food of kharza are encountered: young wapiti, spotted 
deer, roe deer, goral, squirrel, 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). Car- 
rion (except fish) is not taken by kharza and it rarely returns to its 
prey. Captures of sable are a frequent occurrence, increasing with 
increase in [sable] numbers. It does not attack domestic fowl, but 
the old toothless individuals sometimes feed on discards. 

Judging from the fact that kharza always possesses fat depos- 
its, 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 lacking. 

Daily activity and behavior. There is no information for the 
USSR. In India the kharza hunts by day (Pocock, 1941). It travels 
in pairs, but in winter, litters not yet dispersed of up to 5-7 indi- 
viduals are typical. The kharza hunts hoofed animals (musk deer, 
roe deer) in groups surrounding the prey, as observed in India. In 
this way, the yield of the hunt is increased, and the hunting of this 
animal is determined by whether they are in pairs or a group. 
When hunting ungulates, it often tries to drive them onto a smooth 
ice surface (Yu.A. Salmin and V.D. Shamykin). In March, with weakly 
crusted snow, the kharza overtakes a musk deer after 800-1000 m, 
since the weight load of the musk deer on 1 cm'^ of surface is equal 
to 80 gm, and of kharza — only 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. 

The kharza hunts mainly on the ground surface, but climbs 

614 trees proficiently. In 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 
easily tamed. It is active both day and night (Bromlei, 1956). 

Seasonal migrations and transgressions. After March snow- 
falls, when snow depth reaches 50-60 cm, the kharza sinks into 
the snow and prefers to move about in the tree tops, gradually 
descending to the montane foothills to broad-leaf forests in places 
with little 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 May. Litters 
contain two-three young rarely four. Young have not been ob- 
served in summer (Bromlei, 1956). 

Growth, development, and molt. There is almost no informa- 
tion. 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 popu- 
lation 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 deer. The kharza population began to increase only in 1938, 
after the prohibition of 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.). 

Practical Significance 

The value of kharza as a furbearer is not great and does not com- 
pensate for the effort expended in hunting and capturing it. In the 
hunting industry, the kharza causes significant harm by destroying 
a valuable furbearer — sable. In the light of recent data on the diet 
of kharza, this harm is, possibly, somewhat exaggerated. It is de- 
sirable that the number of kharza be limited, but by no means 
achieving complete destruction of this animal, which is 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 42- 
110 (P.Yu.). 

Genus of Wolverines 

Genus Gulo Storr, 1780 

1775. Gulo. Frisch Natur.-Syst. d. vierfiiss, Thiere, p. 17. The 
International Commission on Nomenclature, has decided that 
Frisch' s book does not satisfy nomenclatoral requirements 
and the 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, p. 25 Gulo sibiricus 
Pallas = Ursus gulo Linnaeus (V.H.). 
Dimensions large. 

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 (its width 
less than 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 
auditory bullae relatively small, flattened, thick-walled and wid- 
ened in transverse direction. 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 1 
I-C-P-M- = 38. 
3 14 2 

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 
powerful than those of badger). 

Upper molar typical marten-like in structure — elongated in 
transverse direction, with its inner blade a little wider than outer, 
but small. Upper carnassial tooth very large and strong (its longi- 
tudinal diameter exceeds that of the upper molar, by nearly four 
times), but the inner cusp in its anterior part is relatively weak. 
The main apex of this tooth is high and massive, with sharp lateral 
cutting edges. Lower carnassial tooth large, with two massive apices 
of almost equal height. 

First premolars of both jaws very small and somewhat crowded 
into toothrow, 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 
616 of the upper carnassial tooth is parallel or almost parallel to the 
longitudinal axis of the skull — in all other genera represented in 
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 form of a fist, slightly divided 
anteriorly. 

'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. 
Facial part of head somewhat extended, eyes relatively small, ears 
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 charac- 
ter 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 
to hold only 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 sys- 
tematic relationships of the family as they are described at the 
present time. Among the real features of the genus, it is not pos- 
sible to distinguish those which, against the background of all 
genera, could have supergeneric significance. 

Concerning systematic position, the genus is definitely closer 
to the true martens of the genus Martes. They are closely related 
to 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. 
Only some, actually secondary features in the general 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 all species 
of the family, it 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 its 
recent shape — taiga and forest-tundra, and in part, even tundra. 
618 Quaternary wolverines have been derived from the lower Pleistocene 
G. schlosseri and they do not differ from recent wolverines as 
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 con- 
sider 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 the taiga and a considerable part of the 
tundra zones. 

As a fur-bearing animal, of secondary importance, but is in 
part injurious to the hunting economy (V.H.). 



♦Parentheses omitted in Russian original — Sci. Ed. 

'The placement of the Holarctic wolverines in one species was recently convinc- 
ingly 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. 14, p. 35, tab. 2. Upper 

Ob' river. Requires confirmation — Altai, 
1792. Ursus gulo albus. Kerr. Anim. Kingd. Syst. Cat. No. 381, 

190. Kamchatka. 
1816. Gulo vulgaris. Oken. Lehrb. Naturg. 3, 2, p. 1004. Renam- 
ing of Gulo. 
1820. Gulo borealis. Nilsson. Skand. Fauna. Dagg. Djur. 1, p. 95. 

Renaming of Gulo. 
1829. Gulo arcticus. Desmarest. Mammalogie, 174. Renaming of 

Gulo. 
1829. Gulo arctos. Каир. Entw. Gesch. Nat. Syst. Europ. Thierw. 

1, p. 68. Renaming of Gulo. 
1918. Gulo biedermanni. Matschie. Sitz-Ber. Ges. naturf. Freunde. 

Berlin, p. 147. Mountains south of Telets Lake, Altai. 
1918. Gulo wachei. Matschie. Ibidem, p. 147. Sources of Katun' 

river north of Belukha, Altai. 
1922. Gulo kamtschaticus. Dybowski. Arch. Tow. Nauk. Lwow, 1, 

p. 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 this animal [rosomakha] actually somewhat unusual, is 
written with a double "S" [in Russian]. At the same time, this word has nothing at 
all to 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 unbut- 
toned" (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 Scan- 
dinavia 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 simi- 
lar 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 

appearance, short and broad, especially the posterior part of the 
trunk, on strong short legs and very broad feet. Tail short, approxi- 
mately equal to length of head. It is clothed in long shaggy hairs, 
weakly demarcated from the croup which is covered by dense long 
fur. In its 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 conveys 
the impression of being heavy and clumsy. It usually moves in 
jumps, somewhat laterally and as if stooping — the posterior part of 
the trunk is higher than the shoulder. Although all of its 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 sur- 
face of the feet are easily seen. Claws are light-horn, lighter at 
their tips; on the foreleg their length is 24-26 mm, on the hind — 
22-24 mm. 

Pelage of the wolverine in winter is dense and long, not com- 
pactly arranged but "shaggy" and harsh. The underfur is not very 
dense, length 30-35 mm 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 — 120-150 mm. Therefore, the animal's trunk especially 
posteriorly, seems to be enveloped by a band ("skirt") of long 
hanging, undulating hair, imparting to the trunk in no little degree 
its peculiar appearance. On the legs, 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 
mm. Therefore, the tail is very thick and fluffy, although short. 
620 The fur of wolverine is of low value due to its coarseness, but 
it is warm, sheds water well and is very durable — one of the most 
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 fur. There, much money is paid for skins of this species. This peculi- 
arity of wolverine fur was confirmed by investigations of the American military de- 
partment, who use it for special uniform parts in the Arctic (Krott, 1959). 



928 

Spring molt is complete, but in autumn probably, there is no 
shedding (completion of hair growth). In the first autumn, the animal 
is fully covered with adult pelage. 

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 
zygomatic region, is situated a lighter, sometimes whitish or yel- 
lowish-white field, due to the presence of light ends of guard hairs. 
Ears are dark-brown, slightly lighter on the inner side. The occipi- 
tal 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 
dark-brown ends, the deep underfur is brownish-gray, with ashy 
tinges. The general tone of this "saddle" is 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 out- 
lines 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 — they are darker than the saddle and the legs 
are near-black or black. The tail, except of the above-mentioned 
part, 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 
breast 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. 



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, — deviations from the type de- 
scribed [above] are very great. Very dark wolverines (dark-brown 
in the terminology of furriers) have a very dark dark-brown saddle 
tone, and all remaining body parts are darker than described. The 
breast-band begins farther back, is narrow, and does not include, 
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 field is weakly de- 
fined or almost unnoticeable. 

Wolverines of the light color type ("light-brown") have the 
saddle of light-brown color, and general tone of the remaining 
dark part of the skin is correspondingly light. The breast-band is 
well defined, distinctly observable on the broad shoulder blade, 
particularly posteriorly, and covers a considerable part of the tail. 
621 It is light-ocherous or even whitish. The frontal transverse band is 
sharply defined and very light. In animals of this type, light fields 
are generally no less developed than dark (not considering 
abdomen). 

In the extreme cases, the light parts of the skin occupy a very 
large area, and have sharp outlines and very light straw-yellow 
color. The dark areas are greatly reduced in extent and color — they 
are light, reddish-brown or brownish-red. Starting 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 
ring. Since the stripes of the breast-band are very broad, the saddle 
itself is represented only as a small rounded dark field in the middle 
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. Body length of our wolverine (males and females) 
is 70-105 cm; tail 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. 

Condylobasal length of the skull (old and adult, i.e. older than 
1.5 years) of males is 132.7-158.0 mm, of females, 128.7-149.0 



930 

mm; zygomatic width of males, 92.7-110.0 mm, of females, 89.0- 
103.0 mm; 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, 
of females, 49.0-56.8 mm. Weight in winter time is 10.7-19 kg 
(after Stroganov, 1962, supplemented by material from Z[oologicaI] 
M[useum of] M[oscow] U[niversity] and other sources; total about 
100 individuals)'. 

The measurements are subject to quite significant sexual vari- 
ation — females are smaller and lighter than males, with this differ- 
ence being perhaps quite significant. Some geographic variation in 
dimensions is present (see below). 



Morphological characteristics of the wolverine are explained 
by Krott (1959) in his monograph of this species, by the fact that 
in the past as well as at present, the animal is associated with bogs. 
This opinion is incorrect and is explained by the fact that the 
author's own observations, carried out in a very limited territory in 
the northern part of the Scandinavian Peninsula, were mechani- 
cally 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 a strong and ancient connection with snow. Also 
incorrect is the idea of the cited author, that the connection of this 
animal with bogs itself represents a known sort of biological 
vicariance with the wolf, which is believed to represent its antago- 
nist. In actuality, the picture is quite the opposite — carcasses of 
animals rent by wolves (moose, reindeer) facilitate the winter life 
of wolverine. A connection of wolverine with reindeer has been 
noted thusly; in several places it vanishes together with the disap- 
pearance of reindeer (V.H.). 



"Information given in the monograph of Krott (1959); weight — 20-25 kg, indi- 
vidual males up to 35 kg, females not 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. 



Systematic Position 



931 



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 
in part the tundras of the Old and New Worlds. 

Geographic Range in the Soviet Union 

This is very extensive and constitutes more than half the range of 
the species and almost all 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 is, in many 
cases, difficult since the wolverine 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 wol- 
verine 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 the range given below, if not otherwise 
stipulated, are data 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 is, apparently, connected with the krummholz 
belt 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, 
passing along the Malygin Strait — (Zhitkov, 1913), therefore reach- 
ing approximately 73° N. lat. This is the extreme limit of its 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 
very far to the north of the peninsula (north of Taimyr Lake) and 
nearly reaches its northern tip, probably including it entirely. The 
northernmost occurrences of wolverine in Taimyr are: lower Kheta 
and Khatanga (72° N. lat. frequently); Omulev Bay in the Yenisei 
Gulf at 7240' N. lat. mouth of the Khur, left tributary of the 
Pyasina (73° N. lat.); Novaya and Bol'shaya Balakhnya rivers (about 
73°20' 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 
coast northwards to the latitude of the Faddei islands, i.e. to 77° N. 
lat. or a bit to the south (E.I. Shereshevskii), and the northern 
extremity of Chelyuskin Peninsula (Rutilevskii, 1939). This is the 
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 New Siberian islands — from the 
Bol'shoi Lyakhovsk Island (E.I. Shereshevskii) and even Novaya 
Siberia (Tugarinov, Smirnov and Ivanov, 1934). On the islands, 
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 Kolyma. In all 
events, in this region, the wolverine lives along Omolon, and the 
625 Bol'shoi and Malyi Anyui. Beyond, the range extends over the 
whole basin of the Anadyr' to its mouth, including its northern 
tributaries, and, apparently, at least to the 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. 
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 distribution of the wolverine in the south 
of the Far East is poorly revealed (information is partly contradic- 
tory; Ognev, 1935) and, apparently has a quite odd form. In Ussuri 
Territory, the range occupies the Sikhote-Alin' [range], descend- 
ing along it approximately to 44° N. lat. (in Sudzukhinsk preserve, 
it is absent; G.F. Bromlei). This is 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, appar- 
ently, somewhere in the region of the mouth of the Gorin. From 
here, it directs itself westward, across the upper parts of the Kura, 
Urmya and Bira, 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 
Lake, the southern border of wolverine range in the USSR passes 
outside of the 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 
Lake, 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 53°30' 
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. 



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° paral- 
lel, it abruptly turns again to the west, going along the Kama and 
Volga, seemingly to around the mouth of the Oka and farther, 
probably, along the more northerly part of the Oka- Volga interfluve, 
passing north of Moscow. It is most likely that, a very long time 
ago, it 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 et al., 1955), 
i.e. not south of the Volga. It may be assumed that throughout the 
forests in the region between Moscow and the Urals, the range was 
local, in its southward extent, for example, along the Sura. 
626 West of the Moscow meridian, the range descended signifi- 
cantly southward. It occupied Smolensk district, apparently, to its 
extreme southwestern parts, and probably included the extreme 
western parts of Bryansk district also (Klintsov region), Byelorussia 
and all the Baltic republics, including Lithuania and, probably, 
Kaliningrad district. The most southern place of occurrence in the 
west was along the right bank of the Dnepr — Belovezhsk Forest 
(Brinken, 1828) to the Pripyat' (Pinsk woodland), the Ovruch 
region (former Ovruchsk county), the Pripyat'-Usha interfluve 
(flowing into the Pripyat' not far from its mouth), west of Kiev 
near Radomyshr (border of former Kiev and Radomyshl' coun- 
ties), 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 less permanent residence of 
the wolverine in the 7th cent, (about 49° N. lat.). Farther south, 
only transgressions were known at the end of the 18th century to 
Davydon Brod on the Ingulets river (100 km northeast of Nikolaev; 
about 47°15' N. lat.). This is the most southern point in the Euro- 
pean part of our country where the wolverine was recorded. 



937 

In the past, the wolverine was not encountered along the left 
bank of the Dnepr south of Chernigov ("Chernigovsk 
woodlands" — the northern forest parts of the former Chernigovsk 
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 — although the range never reaches 
the coast — constitute the western border of the range. 

Therefore, at the southern limits of its distribution, the wolver- 
ine was also found in the forest-steppe and transgressions into the 
steppe zone occurred (where it still occurs at present — see below). 

The range of the wolverine, and especially its southern border, 
was subjected to quite significant, and in some places very great, 
changes, during the last century. This is related to both the direct 
extermination and exclusion of the animal, but chiefly to the fell- 
ing and thinning of the forests. At the present time the wolverine, 
which was also previously found in regions of broad-leaf oak-dark 
coniferous forests, is now restricted in its distribution to taiga 
coniferous forests. Therefore, the southern boundary of the range 
in the European part of the Union retreated northward very 
strongly — 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, associ- 
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. Concerning the distribution of the animal in the Altai and to 
its north, there are no data. In western Siberia, the southern border 
of more or less permanent occurrence of the wolverine in the 50' s 
passed, apparently, along the line: Tomsk-Kozhevnikovo-Baksa river 
(southwest of Tomsk — upper Om' — divide between the Tartas and 

■Places of occurrence after S.V. Kirikov (1952, 1959, 1960). On S.V. Kirikov's 
map (1960), instead of 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 
since in the text there is no reference of this sort. 



938 

Тага (data of Stroganov, 1962) — Tara-Tobolsk-Irbit. In the Urals, 
the southern border of the range passes along the line: Irbit- 
Egorshino-Perm' (Shvarts, Pavlinin, Danilov, 1951). Farther on, it 
627 goes a little north of Kirov, again ascends northward in the Nikol'sk 
region (to the Yug river south of Velikii Ustyug), goes farther to 
Tot' ma (at 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 men- 
tioned 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 disap- 
pearance 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 in our, century, individual obser- 
vations of wolverine were known south of the described line. On 
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- 
sent occasional long-distance transgressions of single animals. They 
reach not only the southern border of forests, but also the forest- 
steppe 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 
century, semidesert) in the area north of Lake Ghana (Ghanov, 
Kuibyshev and Vengerovsk regions of Novosibirsk district, 1948- 
1951), 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 

'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 district (1943; Puzanov et al., 1955), in different locali- 
ties in Yaroslavl' district, in particular the Lyubim region, in 
Novgorod district (1925), to 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, 
1957), to the Slutsk region in Byelorussia (see above), in the north- 
ern parts of Kiev district and to Kiev, in Radomyshlya (1914) and 
Proskurov (1924; Sharleman', 1952)* regions and a series of others. 

Geographic Range outside the Soviet Union (see map in genus 
description) 

In North 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 
(the northernmost point of the species range as a whole — north of 
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 New Brunswick, Maine 
Vermont, 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. From here and 
as far as the Pacific Ocean, the range passes far to the south in the 
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 Pa- 
cific Ocean. On the Atlantic coast, the range occupies Newfound- 
land, 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 
Scandinavian Peninsula (in the extreme south, transgressions oc- 
cur in recent times), Finland (at present, absent in the southwestern 
part), and central Europe to Braunschweig and Saxony and to Po- 
land (in the mainland of Europe, it has been absent for a long 
time). In Asia, outside the limits of USSR, the range extends south- 
wards to the northern part of the Mongolian Republic. In north- 
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 47°45' 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 was encountered in Poltav and Voronezh districts, in the Crimea, 
in the Caucasus (Trans-Caucasus), in Hungary, Czechoslovakia, 
Poland; German Democratic Republic and Federal Republic of Ger- 
many, Italy, Belgium and England, France, Switzerland, Austria, 
and Rumania (Pidoplichko, 1951; Krott, 1959) (V.H.). 



Geographic Variation 

The wolverine is referred to as one of the few mammalian species 
in which, with a vast range, reveal insignificant geographic vari- 
ation. This variation bears a clinal character, and in the 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. Geographic variation of this character is expressed in 
the lightening of color from west to east. This lightening occurs, 
however, not by means of gradual lightening from one population 
to another, but through changes in relative constituents of the sepa- 
rate color types described above, and of intermediate between them. 
In this way, wolverines in the west are darker — "dark-brownish" 
prevails, while others are found in which the breast-band is almost 
undeveloped and extremely light i.e. "light-brownish" are few; those 
in which the saddle is in the form of a small spot are completely 
absent or extremely rare. In 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 — interme- 
diate — 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 in population. 
Breast-band not broad, dark or moderate color intensity. Its ante- 
rior ends do not unite on shoulders. Light individuals absent or 
rare. 

Condylobasal length of male skull 132.7-151.3 mm, of fe- 
males, 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). 

European part of country and in West Siberia. Borders in 
relation to following form unknown. 

Outside the USSR — Scandinavian peninsula. 

2. East Siberian wolverine, G. g. sibiricus Pallas, 1780 (syn. 
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 in relation to both other forms not clear. 
629 Outside the USSR — in the northern part of Mongolian Repub- 
lic and 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 doubt- 
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 
belongs in the synonymy of the nominal race, since it is possible 
that it, and not the eastern form, lives in Altai. In order not to 
create new names, it probably is rational to transpose the type 
locality of the form sibiricus 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. 
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 straw- 
yellow 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 mm, of females, 54.2-56.8 mm (ZMMU and Stroganov, 
1956). 

Found in Kamchatka and extreme northeastern Siberia (Anadyi 
Territory, 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. 



In America until recently, the existence of one wolverine spe- 
cies with four subspecies has been accepted (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 

It is primarily that the American form, craniologically well 
differentiated from the contemporary European one, is very simi- 
lar, apparently identical, with the form of the European Pleistocene. 
Evidently, the rate of evolution of the Eurasian wolverine was 
more intensive than that of the American (V.H.). 

Biology 

Population. The wolverine is a rare animal, never forming high 
densities anywhere. Indications of its relative numbers are sup- 
plied by information on prepared skins of hides: 70% of the catch — 
Siberia and Far East, 20% — Urals and 10% — European North. 

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 ungu- 
630 lates. 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 
hectares) (Krott, 1959). 

On the basis of approximate calculations of density and areas, 
the number of wolverines in the USSR may be determined as 7,000- 
7,500 individuals. 

Habitat. The wolverine is widely distributed within the con- 
fines of coniferous taiga and forest-tundra. It also enters the open 
tundra, but here it is rare. It much more rarely penetrates into the 
zone of the mixed forests and to the West Siberian forest-steppe. 

It is encountered both in the plains and in montane conditions. 
Toward its landscape surroundings, it is quite undemanding, 
although it has been noted that it is somewhat attracted to marshy 
areas (Krott, 1959). Deep snow cover usually provides the wolver- 
ine with an advantage in following its prey, and, therefore, the 
main part of its range lies within the boundaries of deep snow; 50- 
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) may also be considered a favorable factor (Krott, 1959). 



944 

In summer, a time of more food security, the wolverine lives 
in very different habitats. Before the young become independent, 
the females are, probably, considerably less mobile than in winter 
and do not go far from the den. 

In winter, and in the period of calving of ungulates, the wol- 
verine confines itself to the region of their concentration. In Altai, 
wolverine avoids human settlements. In Lapland, however, dens 
with litters located 2.5 km from a village have been noted (Krott, 
1959). 

Food. The food composition of wolverine is essentially differ- 
ent in snowy 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 1948- 
1956, 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. 

The larvae of wasps are of great importance in the nutrition of 
wolverines 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 pre- 
fers 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, am- 
phibians, reptiles and crawfishes have not been noted; however, on 
the Pechora, it feeds on the carcasses of spawned-out salmonids 
cast up on the bank. It is only rarely that, in the snowless period, 
it 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 is fairly well studied. Most frequently, these 
are reindeer and elk. In the montane taiga of Siberia, evidently 
they locally 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 
occurrences. In Pechora preserve (Teplov, 1955), with elk and 
reindeer numerous, feeding on live animals is much more frequent. 
631 All ungulates in the food of wolverine constitute from 45 to 75% 
of occurrences. Everywhere, the victims of wolverine are, first of 
all, those weakened from various causes, diseased and injured 
animals as well as juveniles. Reindeer become its victims notably 
more often than elk. Among the latter, the young more frequently 
die (Teplov, 1955). The share of carrion sharply increases with the 
appearance in 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 
event when the wolverine attacks as prey the weaker predators — 
otters, pine martens and foxes. The latter are most often attacked 
in lairs while sleeping, sometimes near carrion. Not rarely, wolver- 
ine 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). 

Wolverine, systematically though not often, rob the traps of 
hunters eating the bait and the catch. When possible, the wolverine 
plunders stocks in the hunter's hut. In rare cases, it 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 to 2 kg of meat, consti- 
tuting about 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 maximum volume of its stomach (Teplov, 
1955). Therefore, current opinion about the voracity of this animal 
is not justified. Even with abundant food, the wolverine does not 
eat "to satiety", 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 



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 range. Observations in nature during the course of sev- 
eral years 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 is a settled animal. In Swedish Lapland, one male lived 
within its hunting territory for a period of not less than six years. 

The area of such a hunting territory in the places with little 
food 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 
tetraonid birds in Finnish Lapland, the hunting territories of wol- 
verines were smaller — about 300 km^ for females and maximum 
distance between brood dens — an average of about 20 km. In 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 is very 

632 stable — 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 is smoothed 
out. In several days large carrion sometimes attracts to it several 
wolverines (for example, 3 wolverines in 25 km^). It is possible 
that this occurs among animals from one litter (cases were noted 
when the litter remains with the mother for the winter) or a group 
forming a structural population unit (1 male, 2-3 females). 



947 



Using for the most part one and the same routes, often by its 
own old tracks, the wolverine periodically visits the particular parts 
of its territory. In winter, these are mainly places of elk concentra- 
tion, the winter pastures of reindeer, etc. 

Burrows and shelters. In winter, the wolverine has no perma- 
nent shelter. Its lairs usually occur under the protection of the 
spruce crowns bent down low, and more rarely on hidden elevated 
places. The animal prefers for its lair the places without an open 
approach. On the Pechora, in the severe frosts, the wolverine hides 
in 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 logs in a snow depth of about 1.5 m. 
Sometimes, in such a den, it makes a deep path in the snow, in 
other cases, the lair is protected from above only by the trunk of 
the fallen tree. The nest itself may be lined with dry fur and grass 
or branches of spruce and fir. In montane localities or on the 
seashore, wolverine makes the brood den usually in a crevice among 
rocks or in the small caves in the very isolated places which are 
difficult to reach. In April 1951, a litter of three young wolverines 
was found in Norwegian Lapland, in a snow den. The nest cham- 
ber was located 20 m from the entrance to the den. 

Daily activity and behavior. In the upper Pechora, the wolver- 
ine is mainly a nocturnal animal. Only in spring (March-April), 
i.e. in the period of nursing the young, diurnal activity is observed 
more often — up to 75% of observations — while in the winter 
months, 25% (Teplov, 1955). 

In winter, in Lapland preserve, it is active in the period of 
polar night mainly during the day (Nasimovich, 1948); the obser- 
vations carried out for several years in the Scandinavian Peninsula 
(Krott, 1959) led to the conclusion that a definite diel rhythm of 
activity for the wolverine is absent. 

Wolverine obtains its food by thoroughly concealing itself from 
its victims, by extended pursuit, by ambush (sometimes from tree 
branches), taking away the prey of other predators, eating the re- 
mains of their meals, and carrion. It waits for hare on the paths and 
attacks tetraonid birds while they are resting, in mating places and 
in snow holes. In searching for food, wolverine seeks out the tracks 
of fox, lynx and others, but it 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 it 
conducts its movements in a straightline fashion, mainly between 
concentrations of ungulates. These movements constitute up to 54% 
of its daily activity. 

In searching for food and while fattening, its path is usually 
twisting. 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 move- 
ments on deep soft snow. In Sweden and Finnish Lapland, wolver- 
ines usually periodically examine their vast hunting territory covering 
a circle of about 1 km diameter. The outlines of the daily move- 
ment in these places usually have a serpentine appearance, with 
633 wide loops. Tracking a wolverine there is very difficult, particu- 
larly if it is followed, since it prefers to walk through "strong", 
difficult of access, places (Krott, 1959). 

The wolverine is a very active animal and has great endurance. 
During a winter diel period it moves from 8 to 45 km, averaging 
21 km, but this 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, 
averaging 22 gm, and according to other data (Dul'keit, 1953), 
27-35 gm. 

It climbs trees quite well, sometimes descending head down- 
wards, but it never jumps from one tree to another (Krott, 1959). 
Unsuccessful hunts constitute a common phenomenon for wolver- 
ine, especially when pursuing ungulates. 

Seasonal migrations and transgressions. Regular migrations 
are unknown in the wolverine. Rare cases of the wolverine's 
appearance in 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, may in 
some measure be related to its 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). 

Reproduction. In Pechora preserve, signs of estrus and mating 
do not occur in the period from October to May. Apparently, they 
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 
(Nasimovich, 1948) — in April and the beginning of May. There- 
fore, 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, 
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, is physi- 
ologically impossible. 

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 May 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, 
a litter found in February was demonstrated (Boehm, 1953). Ac- 
cording to data from 13 litters from Sweden and Finland, within 
February-March, the age of the separate litters may differ by sev- 
eral weeks. The earliest case of parturition was on 11 February 
(Krott, 1959). 

A female having offspring remains barren the following year; 

i.e. barrenness in adult females is not less than 50%. This also 

634 occurred in cases when hunters take the litter from her. At Pechora, 



950 

the percentage of barren females is more than 60%. In Copenha- 
gen 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 
cases, in 10 there were two young each, and in three — 3 cubs each. 
Out of 38 cases, litters of three occurred 17 times, two — 18 times, 
four — 2 times, and one — 1 time (Krott, 1959). Average litter size 
in the Scandinavian Peninsula is 2.5. In Pechora, according to data 
from 7 litters, most often 2 young occur (Teplov, 1955). 

In Alaska, judging from limited material (7 females), wolver- 
ine fertility was apparently higher, there were four corpora lutea in 
six cases and 5 in one. Parallel to them, there were 4 and 3 pla- 
cental 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 fe- 
males, but without subsequent pregnancy. Complete sexual matu- 
rity appears simultaneously in both sexes at the age of four years 
(Krott, 1959). In Alaska, sexual maturity of wolverine begins, 
apparently, at the age of two years. Among immature males, it was 
impossible to distinguish morphologically more than one age class. 
They are easily differentiated from those sexually mature by the 
dimensions of the baculum (in 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 
wolverines take place very quickly — in the course of the first year 
of life (Wright and Rausch, 1955). 

Data on the sex ratio in the population and the rate of mortal- 
ity in it are almost absent, and, hence, there are no data on its 
annual growth rate. In Alaska, one female, in February, had among 
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 
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 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) — 83 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 
den with a litter, who, apparently, took part in the feeding of the 
young. On the Scandinavian Peninsula, no signs of male participa- 
tion in the nourishment of young were observed (Krott, 1959). In 
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, from birth 
of the young to their attainment of sexual maturity, it is proposed 
to distinguish 5 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. 
They drink their mother's milk exclusively. 
635 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 re- 
placed by the second — a dark-brownish color. 

The third phase includes age of 10-24 weeks. At the age of 11 
weeks, average male weight is 3.8 kg, of female, 3.5 kg. In this 
period, the second juvenile fur is replaced by the summer fur, but 
differs 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 
themselves. 

The fifth phase lasts from the age of 12 months to the attain- 
ment of sexual maturity. From the middle of April, the young 



952 

wolverines already have acquired their second summer fur, not 
differing from that of adults, and in mid- August of the second year 
of life, the winter fur of the adult animal. Summer marks the first 
cases of preying on hazelhens, but usually the wolverine's atten- 
tion is not attracted by tetraonid birds. They make 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 
a young experimental male made an attempt to attack a small elk. 
In July, one in the 3rd year of life killed six sheep in one night. 

Enemies, diseases, parasites, mortality, competitors, and popu- 
lation dynamics. With the exception of man and wolves, the wol- 
verine 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 danger- 
ous 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 
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 com- 
petitors. In all events, with the presence of wolves in the region, 
however, their feeding regime is obviously improved and second- 
ary and occasional food use decreases. 

The diseases and parasites of wolverine have been poorly stud- 
ied. In the upper Pechora, 21% of the wolverines caught were 
infected with nematodes. A significant number of the latter (47 to 
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 
dimensions and its generally oval outline, it is reminiscent of the 
broad palm of an adult human. 

The prints, which usually follow one another are almost al- 
ways well distinguished because, even on the soft snow, the ani- 
mal 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. On one 
hand, it is a fur-bearing animal that provides a durable, warm and 
quite beautiful skin, though not expensive, and it sanitizes the 
taiga and tundra, destroying carrion and diseased and dying ungu- 
lates. 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 danger- 
ous. In Sweden, the wolverine is considered a useful animal for the 
forest economy since it reduces in the number of ungulates in areas 
impossible for hunters to reach. The number of ungulates is, in many 
cases, very great 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 destruc- 
tion the year round. In practice, this is difficult to realize. In the 
State preserves, if its numbers increase significantly, it must be 
limited. 

The world catch of wolverine comprises about 6,000 skins. Its 
relative contribution in the 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 
particular jawed traps, take them poorly. Setting the traps around 
carrion give somewhat better results. For hunting with dogs, fero- 
cious and strong laika dogs are required, since the wolverine fiercely 
defends itself. Wolverine is very resistant to the action of poisons. 



954 

All methods for its capture are complicated by the vast area of its 
home range and its very long daily movement. 

In the Scandinavian Peninsula, the best effect is attained by 
searching out litters, which is carried out in the second half of 
March and the first half of April. Two or three hunters on skis 
with a 2-3-week stock of food search a typical area for fresh 
tracks of a female leading to a den. In March, when the young are 
still blind, the female almost 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 fe- 
male — 70. Moreover, the hunters receive a bounty: 80 dollars for an 
adult wolverine and 40 for young. 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 d. vierfUss. Thiere, p. 11. 
1817. Putorius. Cuvier. Regne anim., 1, p. 147. Mustela putorius 

Linnaeus. 
1829. Arctogale. Каир. Entw. Gesch. Nat. Syst. Europ. Thierw. 1, 

p. 30. Mustela erminea Linnaeus. 
1829. Ictis. Каир. Ibidem. 1, p. 35, 40, 41. Mustela vulgaris 

Erxleben = Mustela nivalis Linnaeus. 

1840. Foetorius. Keyserling et Blasius. Wirbelthiere Europas. p. 
68. Mustela putorius Linnaeus. 

1841. Gale. Wagner. Schreb. Saugeth. Suppl. 2, p. 234. Mustela 
vulgaris Erxleben = Mustela nivalis Linnaeus. 

1841. Lutreola. Wagner. Ibidem. 2, p. 239. Viverra lutreola Linnaeus. 

637 1871. Mustelina Bogdanov. Birds and animals of the chernozem 

belt of Povolozh'e (Tr. Obshch. estestvoisp. pri. Imp. Kazansk. 

Universit., 1, 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, p. 167. Mustela lutreola 

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, slightly widened in 
the occipital region, the facial part is 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 or very weak or else the postorbital process is barely de- 
fined. The dorsal profile of the skull is 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 de- 
pressed. The diameter of the infraorbital foramena is greater than 
that of the alveolus of the upper canines; in some species, they are 
equal to it or even less. 

In a few forms, the whole skull is wider and shorter, the brain- 
case is relatively shortened and the facial part is strengthened, the 
zygomatic arches are relatively robust and more widely separated, 
the postorbital constriction is relatively well or even strongly 
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. 

The auditory bullae in all species are notably swollen through- 
out the lower surface, usually of elongated bean-like form, and 
situated more or less parallel to each other; in some species, they 
are relatively shortened and the posterior portions are relatively 
more widely divergent than the anterior. The hook-like processes 
of the pterygoid bones are not united with the auditory bullae. The 
mammary (mastoid) processes are weakly developed, and the lat- 
eral occipital (paroccipital) are not developed. The bony palate is 
of moderate width. In general, with a comparatively large number 



956 

of species, several of them have evident differences in skull struc- 
ture, though in close species, they are not great (see later). 

3 13 1 
Dental formula I-C-P — M — = 34. There is no additional cusp 

on the inner side of the main crest of the last lower premolar (third 
tooth behind canine — ^just anterior to carnassial). The lower car- 
nassial tooth (first molar) has no 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 
it is not lost or this occurs only rarely. 

The trunk is thin and strongly elongated, in some species to an 
extreme degree, and it appears as though "snake-like". The legs 
are short, in several, very short, the tail is of various lengths and 
may attain half or nearly half the body length. In several species 
it is short, and constitutes about 0.25 of the body length or less and 
perhaps equal the length of the hind foot. The head is relatively 
638 small, narrow, elongated and flattened, usually no broader in diam- 
eter than the neck, in some almost "snake-like" in appearance. The 
ears are small, very widely separated, rounded; the eyes are quite 
large. The extremities are digitigrade; palms and soles are moder- 
ately broad. 

Anal scent glands are developed in all species, 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 
length over the whole body. In the majority, the fur is soft and 
silky and in some, highly valued. In the majority, the tail is cov- 
ered by short hairs and is relatively thin, but in some quite fluffy. 

The color is quite variable — two-toned with a light venter, of 
monocolor ocherous or brown tones, or blackish-yellow with a 
dark venter. A distinct throat patch does not occur. In some spe- 
cies, there is a pattern on the head in the 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). 



957 

Seasonal dimorphism in density and length of the fur of north- 
ern forms is very pronounced, while in the south, it 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 
limits of 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 ab- 
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 bur- 
rows. They inhabit very diverse biotopes — from the Arctic and 
high mountains higher than 3000 m above sea level to steppes and 
deserts and tropical forests. All of them can swim — some species 
are typically amphibious forms associated with fresh water (minks). 
Some species do not avoid proximity to humans and even reveal 
some anthropophilous tendencies. 

They are settled forms; monogamous; there are 1-2 and 3-4, 
to 10 young. Lairs are made in extremely varied places — among 
windfalls, in rocks, etc., but chiefly in underground burrows, mainly 
of rodents. They do not 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 distinc- 
tive 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. Some- 
times there is a quite close connection with particular species of 
rodents (the steppe polecat — ground squirrels, ermine — water vole, 
weasel — small voles 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 forms a body length from 130 mm 
(averaging about 160 mm) and a weight of less than 100 gm 

*Lit. "mouse-eaters". 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. 

The range of the genus is very vast and occupies the greater 
part of Eurasia 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 
640 entire Arctic archipelago and the northern and northeastern part 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 Indies. On the islands of the Bering Sea (Aleutians, Pribilofs, 
St. 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 cor- 
ner^ — 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 New Siberian Islands. The southern limit in Asia 
includes, to the south, Palestine, Syria and the northern half of 
Iraq. In Iran, the range occupies the northwestern part of the coun- 
try, 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), whence the border, 
apparently, to the south covers the 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 
Hainan. On the Arctic Islands, except New Siberian, as in the 
Commander Islands, representatives of the 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 some- 
times changed in recent times, either negatively or positively. 

The scope accepted here of the genus (13 species) itself rep- 
resents a quite heterogeneous group. 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 
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 genus accepted here into sepa- 
rate genera or subgenera. Thus, separate groups 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, if we consider the group of species as a 
whole, it is sufficiently clear that strict delimitation of it into sev- 
eral is impossible, and the most extreme forms, as shown above, 
641 are connected with each other by a series of intermediates. This 
applies all the more to other members of the genus. Therefore, it 
is natural to accept the large genus Mustela in 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 



961 



(Stroganov, 1962) cannot be considered well founded if one 
considers all species of the genus. The polecats even now are 
sometimes separated into a separate genus {Putorius), that has been 
poorly received. The mink serves as a connecting link between 
them and the remaining species^. 

The identification of the actual systematic position of the ge- 
nus among these various forms (genera) present in the subfamily, 
is very difficult. It is apparent, however, that this is a line of 
development in the subfamily different from the true martens and 
wolverine. It is more connected with the other genera of the sub- 
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 is indicated by the craniologi- 
cal structure of the species belonging to the subgenus Mustela 
(their specialization obviously proceeded in another direction, in 
particular the elaboration of a special 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 be- 
tween these fossil forms is not clear. 

The following 13 species may be included in the genus (listed 
in order of increasing specialization): weasel, M. nivalis 
(including M. rixosa which American authors consider a separate 
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 



^An indication is that hybrids may exist between the black polecat and the Rus- 
sian mink in nature. For details on relationships in the genus, and connections between 
the polecat group on one hand and a remaining species see section on polecats and mink. 

*Local name referring to suslik, or ground squirrel, which in many places is an 
important prey species — Sci. Ed. 



962 

or steppe polecat, M. eversmanni. The last three species belong to 
the subgenus Putorius, and the remainder — 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. 
nigripes), 2 are European (M. putorius, M. lutreola, the latter also 
in the 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. (Л/.) 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 pole- 
cat, M. (P.) eversmanni Lesson, 1827. One species — the Ameri- 
can 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 Identification of the Species in the Genus Mustela 

1 (4). Color of entire body monotone white (winter) or lower 
side of body pure white (sometimes with light grayish- 
yellow highlights), upper side various brownish or sandy 
tones (summer). Color of upper lip and chin not differen- 
tiated from that of surrounding parts of head. Color 
borders on sides well marked. Skull moderately elongated — 
distance between mastoid (mammary) processes approxi- 
mately equal to half of condylobasal length of the skull. 

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 

ermine, M. (M.) erminea (page 995). 

3(2). Tail monotone along entire length (there may be indi- 
vidual dark hairs at its extreme end). Length of tail 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 to 47.2 mm; of female 
up to 36.7 mm weasel, M. (M.) nivalis (page 965). 

4(1). Color otherwise. If dorsum tawny and brown tones, and 
venter light and border between dorsum and venter colors 
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. 

5(8). Lower surface of body, except throat and chin and some- 
times 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 mas- 
toid 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 its edges form parallel or almost parallel lines 

black polecat, M. (P.) putorius (page 1107). 

7(6). 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 is present in some races of solongoi [M. altaica]; see 14 (13) and 
text. 



964 

8(5). Color of abdomen and lower side of body not black or 
blackish, legs not black, tail without black color, its color 
corresponds to upper body. Skull moderately elongated 
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, M. (M.) lutreola (page 1078). 

11 (10). Only lower lip 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- 
what more mediad to antero-outer corner of carnassial 
tooth American mink, M. (M.) vison (page 1391). 

12 (9). Entire body color if not monotone is not dark tawny- 

brown, but bright yellowish red or dull yellowish-straw- 
sandy, or below is somewhat lighter than above, or vice 
versa. Skull elongated — distance between mastoid processes 
is usually less than half of condylobasal length of skull. 

13 (14). Color of muzzle and area near eyes coffee-brown, form- 

ing well delineated dark "mask" on head. Postorbital con- 
striction of skull 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) 

Siberian weasel, M. (M.) sibirica (Page 1051). 



965 

14(13). Dark "mask" on muzzle absent. Postorbital constriction 
of skull short, its lateral outlines not parallel and forms 
a sharp isthmus directly behind supraorbital processes. 
Color is bright or dull, yellowish-red, lower side a little 
to 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 mm, of females — less than 48 mm) 
solongoi M. (M.) 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. 
1811. Mustela gale. Pallas. Zoogr. Rosso-Asiat. 3, p. 94. Renam- 
ing of vulgaris. 
1877. Mustela stoliczkana. Blanford. J. Asiat. Soc. Bengal, 46, 2, 

p. 260. Yarkand. 
1896. Mustela rixosa Bangs. Proc. Biol. Soc. Washington, 10, p. 

21. Saskachewan, Canada. 
1899. Mustela nikolskii. Semenov. Zap. Akad. nauk, ser. 8, vol. 6, 

p. 14. Vic. Simferopol, Crimea^. 

^^2 ^^^^ correct spelled name for this species is undoubtedly, lazka, and is derived from 

the verb lazat' [=to climb]. The ability of the animal to climb with extraordinary agility, 
especially among fallen limbs etc., is striking, and immediately calls attention to itself. 

^Westerbotten province lies between 60° and 61° N. lat. and therefore, often 
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). 



966 

1900. Putorius nivalis pallidus. Barrett-Hamilton. Ann. Mag. Nat. 
Hist. 5, p. 48. Kokand, Fergana. 

1900. Putorius nivalis caucasicus. Barrett-Hamilton. Ibidem, p. 48. 
Caucasus, Gochal (?) mountains, 12000'. 

1901. Mustela (Ictis) dombrowskii. Matschie. Sitzungsber. Ges. 
naturf.-Freunde Berlin, p. 231. Siul'nita. Romania. 

1903. Putorius (Arctogale) pygmaeus. J. Allen. Bull. Amer. Mus. 

Nat. Hist., 19, p. 176. Gizhiga, northern shore of Okhotsk 

Sea. 
1907. Putorius nivalis dinniki. Satunin. Izv. Kavkazsk. muzeya, 3, 

p. 105. Stavropol (Caucaskii). 
191 1. Ictis boccamela nikolskii. Satunin. Izv. Kavkazskovo muzeya, 

5, p. 257. Crimea^ 

1921. Mustela rixosa namiyei. Kuroda. Journ. Mamm., 2, p. 209, 
Aomori, northern part of Honshu (Hondo) island, Japan. 

1922. Ictis nivalis var. kamtschatica. Dybowski. Arch. Tov. Nauk. 
Lwow, 1, p. 349. Nomen nudum. 

1926. Mustela punctata. Domaniewski. Ann. Mus. Zool. Polonici. 
N.H. 5, p. 55. Darasun; southern Trans-Baikaliya. 

1927. Mustela nivalis mosanensis Mori. J. Chosen. N.H. Soc, 5, 
p. 28. lengan near Mosan, Korean Peninsula. 

1936. Mustela pygmaea yesoidsuna. Kishida. Dobuts Zasshi, 48, 
4, p. 177. Hokkaido, Japan. 

1936. Mustela pygmaea karaftensis. Kishida. Ibidem, p. 177. 
Sakhalin. 

1937. Mustela trettaui. Kleinschmidt. Falco, 33, p. 11. "Germany". 

1952. Mustela nivalis kerulenica. Bannikov. Byull. Mosk. Obsch. 
Isp. Prirody, 57, No. 2, p. 40. Vic. Undurkhan on Kerulen 
[г.], Mongolian Republic. 

1953. Mustela nivalis heptneri. MoToso\a-Turo\a. Zool. Zhurn., 
37, No. 6, p. 1267. Egri-Gek — tributary of Kushka [river], 
Badkhyz; southern Turkmeniya (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 tail not black. Tail length 

■•Satunin, apparently, not knowing of the work of Semenov, describes the Crimean 
weasel as a new form. 



967 

significantly less than half of body length, usually not more than 
one third. Skull is moderately elongated — distance between mas- 
toid (mammary) processes approximately equal to half of condylobasal 
length of skull. Width of skull below canines approximately equal 
to interorbital width. Dimensions small to very small (V.H.). 

645 Description^ 

The general appearance of the weasel is very characteristic. It has 
a thin, greatly elongated, extremely flexible body with a small, 
somewhat elongated head with a small, blunt muzzle. The head is 
narrow, not thicker than the neck; the eyes are quite large, dark 
and slightly bulging; the ears are situated wide apart, short and 
rounded. The legs are short, the feet of moderate breadth. The tail 
is relatively short. It is considerably less than half the body length, 
as a rule not greater than one-third of its length, sometimes not as 
long or slightly longer than the hind foot, and in individual cases, 
even shorter than the hind foot, covered with short, close-lying 
hairs, and appears thin, not 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. Summer, 
coloration is bi-colored — the lower body including the lower jaw 
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 differ- 
ent races, varies greatly — from dark-tawny or dark-chocolate to 
light pale tawny or sandy. The color of the dorsum extends to the 

'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 some- 
times 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 south- 
erly forms do not whiten at all in winter, or only some lightening 
in color may take place, or the side becomes white, but the white 
color does not cover the middle of the back. When molting, the 
animals sometimes become piebald — irregular dark parts are 
scattered over a white field. 

Individual differences in fur color are in general insignificant, 
however, in various races, the amplitude of these differences is 
different and in several (Western Europe), these differences may 
646 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, espe- 
cially the lower jaw. Age and sex differences in color are absent, 
but geographic is, 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 
weak and very short. Width of the skull above the canines is ap- 
proximately 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- 
mately equal to the longitudinal diameter of the canine alveolus. 

The upper profile of the cranial region of the skull is slightly 
flattened and the interorbital area does not form a significant el- 
evation on it. 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 more strongly developed, but is not 
considerable. The auditory capsules are bean-like in form, convex 
and their 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 rela- 
tively 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 por- 
tion, 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 also be expressed. 

Individual variation in the skull is considerable in all charac- 
ters. The sexual is illustrated by the fact that the skull of males is 
larger, and wider in the mastoid part, the postorbital constriction 
is absolutely and relatively narrower, and crests, tubercules etc. of 
the skull are more strongly developed. 

Age variation of the skull is insignificant. In young animals, 
the brain case is still 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 — earlier than in the large members of 
the genus. 

Geographic variation of the skull, in accordance with the gen- 
erally large variation of the species, is very great. It is expressed 
both in general dimensions and also in general structure and pro- 
portions 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 — in the 
smallest forms, it is almost unexpressed, but grows with the in- 
crease in general dimensions and is well developed in the largest 
races. 

The OS penis is sharply bent upwards at its distal end, and here 
is formed a hook, characteristic of the majority of species of this 
genus. 

The dimensions of weasel are subjected in some races to great 
individual variation, rarely found among mammals in general. Thus, 
in the race vulgaris, the weight of the largest individuals may 
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 is 
648 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 
forms 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 vari- 
ation in relative tail length is characteristic — in various races it 
constitutes from 13 to 30% of the body length. In the extreme case, 
in individual animals the tail length may be less than that of the 
hind foot length or equal to it. 

Body length of males is 130-260 mm, of females, 114-204 
mm; tail length of males is 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 
females, 27.5-36.7 mm; mastoid width of males is 11.0-22.4 mm, 
of females, 12.5-18.9 mm; interorbital width of males is 5.0-10.8 



971 



mm, of females, 5.5-7.8 mm; postorbital width of males is 6.8- 
9.0 mm, of females, 6.2-8.0 mm; zygomatic width of males is 
13.8-26.0 mm, of females, 13.8-19.2 mm. Length of the os penis 
is 12-20 mm. 





647 



Fig. 235. Skull of weasel, Mustela (Mustela) nivalis L. 



972 



Weight of males is 36-250 gm, of females, 29.5-117 gm^ 
(V.H.). 

Systematic Position 

Weasel is itself an entirely specialized form; however, within the 
limits of the genus Mustela, it is, apparently, properly placed at the 
beginning of the entire series of species. 

In several respects, it is, compared to all remaining species, 
less specialized. This is shown, first of all, by the above- 
mentioned "infantile" features of its skull. As mentioned before, 
these infantile features are associated, to a considerable • degree, 
with the very small dimensions of the animal, and are particularly 
distinct in the small forms {rixosa group), but they are strong even 
in its large races, which give up little in dimensions to the ermine 
(weasels of the boccamela-heptneri group). 

The weasel stands nearest of all to the ermine (M. erminea), 
although they differ in a series of essential characteristics, includ- 
ing the structure of the os penis. It is probable that this is one of 
the details of the mechanism of reproductive isolation where both 
species are morphologically closer to each other in areas simulta- 
neously inhabited by them (Tien Shan). The weasel is connected 
through the ermine with M. altaica with which it does not differ 
in structure of the genital organs. On the whole, it is quite a close 
group, noteably distanced from the larger species of the genus 
(V.H.). 

Geographic Distribution 

The temperate, and in part, the arctic zone of the New and Old 
Worlds; in the Old World, also in part the subtropical zone. 

Geographic Range in the Soviet 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 Stroganov (1962) and unpublished material 
of L.G. Morozova-Turova; os penis dimensions after Ognev (1935); weight and other 
data also from the collections of the Z[oological] 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 is absent on the Commanders; it is, 
apparently, found on Shantar Islands, although there is no positive 
information about it. It inhabits Sakhalin (information on the ab- 
sence of weasel here is mistaken); of the Kuril islands, it is present 
only on Kunashir; the possibility that it may be met with in some 
of the southern islands is not excluded. In spite of some asser- 
tions, it is also found in the Kyzylkum and Karakum. It is not 
excluded that the weasel does not reach the extreme north of 
Taimyr, and information about its occurrence in the Pamir are not 
completely definite. 

650 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 mountain system. It 
exists in northern Afghanistan, Kashgaria (known to the east as far 
as Lobnor), in Dzhungaria, in the northern part of the Mongolian 
Republic including to the south the Mongolian, and, probably, Gobi 
Altai and Trans-Altai Gobi, Khangai and the Kerulen [river]* re- 
gion, in northeastern China (former Manchuria), on the Korean 
Peninsula and in Japan. In China, it is apparently absent in Inner 
Mongolia and Tibet. The extent of its distribution to the south in 
the eastern parts of China is unknown. It scarcely goes far south, 
although one of the forms was recorded in Tonkin (North Viet- 
nam) 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 in the weasel is not only very great, but also 
complicated and 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 at the beginning of the past century, it was noted that in 
Europe, among large, comparatively long-tailed weasels, there were 
encountered very small short-tailed ones. Later this form received 
the name M. minuta, and began to be considered as a category of 
distinct species, existing in several parts of Europe among 
populations of the larger common weasel — M. nivalis (M. vulgaris). 
Later, it 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 
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 
authors were inclined to consider a separate species (M. 
boccamela). This opinion, on the contrary, had been rejected much 
earlier, but the representation of species independence of the 
European M. minuta was still held by some western authors in the 
40's and even the end of the 50's (Van den Brink, 1958). Ameri- 
can authors, up- till now, consider their weasel an independent 
species — M. rixosa and that the small weasels of Asia and Europe 
belong to its subspecies. 

In actuality, a long time ago it was accepted by our systema- 
tists (Ognev, 1935, for example), that in the whole Palearctic only 
one weasel species exists, geographically very variable, with a 
652 complicated internal structure. Its extreme forms (pygmaea-heptneri) 
are so strongly differentiated that with limited material (territorial 
or quantitative) and insufficiently worked out theoretical 
prerequisites, they are actually very easily accepted as different 
species. Other European investigators gradually came to agree with 
the concept of species unity (Ellermaji and Morrison-Scott, 1951; 
Zimmermann, 1959 and others). It is also doubtless that the wea- 
sels of 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. still demand further 
clarification with large samples. 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 geo- 
graphically: 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 unpub- 
lished 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. 
Outside the USSR — Finland, northern Scandinavian Peninsula, 
Mongolian Republic, northeastern China (former Manchuria), 
Japan, North America. 

Weasels of this group are the smallest of all forms of the 
Carnivora. 

1. Siberian weasel, M. (M.) n. pygmaea J. Allen, 1903 (syn. 
kamtschatica, karaftensis; with respect to the weasel of the Kuril 
Islands, the name namiyei was applied). 

Dimensions very small. 

Tail short, its length equal to hind foot or a bit longer and 
constitutes about 13% of body length. 

In summer coat, dorsal color dark-brown, very rarely lighter, 
reddish. In winter, entirely white. Hair length on sacrum, 10.3 mm 
in 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 foot length, 16-M21-25 mm. 

Condylobasal length of male skull, 22.8-M32.7-34.8 mm; mas- 
toid width, 11.0-M 15.2-16.8 mm; interorbital width, 5.0-M7.2- 
8.2 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), 29.5- 
M41.0-62 gm (material in Zoological Museum of Moscow 
University)'^. 

All of Siberia, except southern and southeastern Trans- 
Baikaliya; northern and middle Urals, in northern Kazakhstan (a 
few south of Syr-Dar'ya mouth), and Far East including Sakhalin 
653 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 
Sakhalin and Kamchatka to Altai. Animals from Perm district, belonging to this 
form or approaching nivalis average, apparently, somewhat larger: males (5) 50- 
M60.8-70 gm. 

* Apparently includes one unsexed animal — Sci. Ed. 



979 

In the European part of the Union, the most typical weasels 
are in the 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 
population is not systematically homogenous and there may be 
distinguished forms here differing from true pygmaea. Thus, it is 
possible that weasels of the southern part of the Far East them- 
selves represent a distinct form (it apparently can be given the 
name mosanensis or namiyei); the position of Sakhalin and Kuril 
weasels is not clear, etc.; West Siberian weasels (except northern) 
are sometimes assigned to nivalis (Stroganov, 1962), etc. 

2. Trans-Baikal weasel, M. (M.) n. punctata Domaniewski, 1926 
(syn. kerulenica). 

According to dimensions and relative tail length (tail consti- 
tutes about 13.9% of body length), it corresponds with M. (M.) 
n. pygmaea, but according to color of summer fur of upper body, 
is somewhat lighter. In winter, it becomes completely or almost 
completely white. Skull somewhat wider in its rostral portion, post- 
orbital 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, 12-Ml 8.2-23 mm'l 

Condylobasal length of male skull (6), 31.0-M32.4-35.0 mm, 
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, 6.6-M7.2-8.5 
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 part of Mongolian Republic. Materials 
in Zoological Museum of Moscow University. 

'''Weight of 21 specimens from collection of Z[oological] M[useum of] M[oscow] 
U[niversity] including individuals, the stx of which 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] 
M[useum of] M[oscow] U[niversity]. 

*Minimum value omitted in Russian original — Sci. Ed. 



980 

This form has its main area of distribution outside the bounda- 
ries of the USSR and reaches us only on the edge of its range. 
Lightening of body color is, apparently, connected with occupation 
of more arid regions. 

B. Subspecies of the group boccamela, large weasels. 

Weasels of very large dimensions, with large skull and rela- 
tively 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 USSR — southern Europe (Mediterranean), Asia 
Minor, Iran, Afghanistan. 
654 3. Trans-Caucasus weasel, M. (M.) n. boccamela Bechstein, 1800. 

Dimensions very large, tail very long, considerably longer than 
hind foot, constituting about 30% of body length. 

In summer fur, color of upper body light brownish or chestnut, 
with yellowish or reddish tints. In some individuals, a brownish 
dot behind the corner of the mouth and sometimes on chest and 
belly. Winter fur not so pure white as in more northern forms; 
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 skull, 41.0-M42.1-46.2 mm; of 
females (6), 29. 8-M3 1.2-36.0 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, 6.0- 
M7. 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 prob- 
ably western parts of Iran. 

The weasel of Trans-Caucasus does not have any essential 
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 large, in general corresponding with preced- 
ing form. Tail very long, considerably longer than hind foot and 
constitutes 25-30% of body length. 

In summer fur, color of upper part of body very light, 
"desertish" — brownish-sandy or pale-yellowish, 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 mm in summer and 7 mm 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 foot, 27-M33.2-35 mm. 

Condylobasal length of male skull, 40. 5-M43. 1-45.5 mm; 
mastoid width, 2 1.0-M2 1.3-22.0 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 Mid- 
dle Asia from Caspian Sea to Semirech'e and southern Tadzhikistan, 
Kopet-Dag. 

Outside the USSR — in Afghanistan and the northeastern part 
of Iran. 

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 meas- 
urements. Sexual dimorphism well developed. 

Middle and southern regions of the European part of the coun- 
try; 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 В 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. 

Color of upper body in summer fur is dark-brownish or chest- 
nut, rarely lighter — yellowish-brownish. Winter fur pure white. The 
hair length on sacrum 9.5 mm in summer, and in winter, 
12.5 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, 22- 
M27.3-30 mm; of females, 19-M22.1-25 mm. 

Condylobasal length of male skull (13), 30.5-M35. 3-40,2 mm; 
of females (9), 29.0-M 32.0-36.8 mm; mastoid width of males, 
15.0-M17. 3-19.6 mm; of females, 12.8-M15.0-17.5 mm; interor- 
bital width of males, 6.0-M7.7-9.5 mm; of females, 5.0-M6.9-7.0 
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 
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 
the middle and southern Urals, northward approximately to the latitude 
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 and even 
Moscow districts, and, at the same time, the weasels of Perm dis- 
trict are already close in dimensions to, or belong to, nivalis. Due 
to this, individual variation of this form is very great, and equally 
so with the relatively large "normal" forms, the extreme small 
variants are found in nearly all of the range of the subspecies and 
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. 



983 

The possibility of accurately diagnosing and designating this 
form is made more difficult by lack of clarity concerning represen- 
tation of the typical nivalis lineage, which, probably, itself 
represents a transition from the northern pygmaea to the 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 in M. (M.) n. nivalis. Tail 
relatively longer, its 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 light- 
brownish to dark-chestnut. Winter fur white and only in extreme 
south of range is whitening incomplete. Hair length on sacrum 
7.5-8.5 mm in summer and in winter 10.5-13.5 mm. Skull larger 
in M. (M.) n. nivalis, and zygomatic arches relatively widely sepa- 
rated. 

Body length of males (26), 173-M2 12.6-243 mm; of females 
656 (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. 

Condylobasal length of skull of males, 31.5-M38. 8-43.0 mm; 
of females (6), 25. 8-M33. 2-36.0 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, 5.0-M8.4-10.2 mm; of 
females, 6.0-M7.1-8.0 mm; zygomatic width of males, 17.0- 
M20.0-22.0 mm. 

Southern regions of the European part of the country from the 
latitude of southern Voronezh and Kursk districts, Crimea, Cis- 
Caucasus, northern slope of the Main Caucasus, eastward at least 
to the Volga (eastern border not precisely known). 

Outside the USSR — Europe southward to the Alps and Pyr- 
enees. 

In some parts of its range in the south, the described form has 
features significantly similar to large weasels of the type boccamela. 
This reveals itself particularly in the northern Caucasus, and is 
partially explained 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 in the accepted sense itself represents a 
transition to the large southern weasels of the type boccamela. In 
the north, the 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 cer- 
tain degree, conditional. 

The majority of western European authors accept for "Ger- 
many" 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 synonym of nivalis, a fact which is un- 
founded. In general, for a correct understanding of the form nivalis, 
a solid revision of the weasels of the Scandinavian Peninsula is 
necessary. Some authors consider the Scandinavian weasel M. (M.) 
n. pygmaea as identical to the Middle European "minuta" and an 
extreme variant ("minus- variant") of Л/. (M.) n. nivalis (Reichstein, 
1957), with which it is difficult to agree. 

7. Montane Turkestan weasel, M. (M.) n. pallida Barrett- 
Hamilton, 1900. 

Dimensions somewhat smaller than in M. (M.) n. vulgaris, 
approximately correspond to dimensions of M. (M.) n. nivalis. Tail 
of moderate length, constituting about 24% of body length. 

Color of summer fur light-brownish. Winter fur white. Skull of 
moderate size, relatively narrow in mastoid part, postorbital con- 
striction relatively short and narrow. 

Body length of males (17), 180-M 192-2 15 mm, tail length, 
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- 
8.4 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 Hindu- 
kush (Afghanistan). 

The scheme presented focuses on the main features and trends 
in geographic variation of the species in our country. It cannot be 
considered sufficiently studied, and is not generally accepted (see 

■"Description of range within USSR omitted in Russian original; this includes 
montane parts of Turkmenia, Uzbekistan, Tadzhikistan, Kazakhstan, and Kirghizia — 
Sci. Ed. 



985 



Ognev, 1935; Novikov, 1956; Stroganov, 1962). The question of 
weasels in the European part of the country and Caucasus is par- 
ticularly complicated. 



657 For the parts of the range lying outside the USSR, the follow- 
ing forms are usually accepted: 1) M. (M.) n. subpalmata Hemprich 
et Ehrenberg — Egypt (Lower); 2) M. (M.) n. numidica Pucheran, 
1855 — Morocco, Algeria, Malta, Azores Islands? Corsica; 3) M. 
(M.) n. stoliczkana Blanford, 1877 — Kashgaria'^; 4) M. (M.) n. 
siberica Barrett-Hamilton, 1900 — Pyrenees [Iberian] Peninsula and 
Balearic Islands; 5) M. (M.) n. galinthias Bate, 1906 — Crete; 6) M. 
(M.) n. russeliana Thomas, 1911 — Sichuan, southern China; 7) M. 
(M.) n. namiyei Kuroda, 1921 — Japanese Islands; 8) M. (M.) n. 
mosanensis Mori, 1927 — Korean Peninsula; 9) M. (M.) n. 
tonkinensis Bjorkegren, 1942 — northern and southern Vietnam (it 
is probable that this race belongs to another species); 10) M. (M.) 
n. alleghaniensis Rhoads, 1901'^ — southeastern part of the range 
in America (Michigan, Pennsylvania, 'Virginia, North Carolina, 
Ohio, Illinois, Wisconsin); 11) M. (M.) n. campestris Jackson, 
1913 — 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 
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 Panfilov — Sci. Ed. 

'*The American authors consider this and the following form as belonging to a 
separate species M. (M.) rixosa. 



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 1 ermine. 

The area of weasel abundance lies more southward than that of 
ermine, and numbers of ermine in southern regions is usually less 
than weasel numbers. This partly depends on differences in possi- 
bilities of capture of mouse-like rodents between these two spe- 
cies. In the European part of the USSR, weasels are found in small 
numbers in the taiga forest zone. In this zone, they mainly live in 
fields 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 
Siberia, it is more numerous in the southern parts. In northeastern 
Siberia, it is rare. It is more often encountered in southern Trans- 
Baikal and Primore Territory. 

Due to significant fluctuations in numbers, moreover not one- 
time, its occurrence in different parts of its range can only be 
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 
658 tracks may change 8-10-fold, but in all cases, in broad-leaf for- 
ests, 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. It is established in tundra, does 
not avoid taiga and is common in the forest-steppe zone and 
chernozem steppes. It lives in grassy steppes, semideserts and moun- 
tains. It does not avoid proximity to humans, settling sometimes in 

Table 62. Frequency of weasel tracks in the broad-leaf [Zhiguli] and 
mixed forests [Kalinin] (along 10 km route) 





Year of high 


numbers 


Year of 


low 


numbers 


Region 




Quantity 








Extreme 


Average 


Extreme 




Average 


Southwestern part, 
Kalinin district 
Zhiguli 


0.7-12.8 
9.7-122.8 


9.7 
36.2 


0.2-8.6 
0-13.3 




0.9 

2.7 








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987 

structures within a town (outskirts of Moscow). Although encoun- 
tered almost everywhere, the weasel is known to prefer certain 
places. Thus, in the middle forest belt, the weasel is most often 
met with in openings, clearings, forest edges, plantations, roadside 
ditches, etc. It is often in riparian vegetation of small rivulets, in 
meadows with haystacks, in mezha*, threshing-floors**, in store- 
houses, barns, and other buildings in villages. In Siberia, the wea- 
sel is more often encountered in steppe than in taiga, where it 
mainly restricts itself to meadows, old burns and in tall grass. In 
the chernozem and steppe belts, it inhabits steppes, fields, pas- 
tures, gardens, vegetable gardens, stonepiles, woodpiles, thresh- 
ing-floor**, hay/grain stacks, fences, inhabited and uninhabited 
structures; and in mountainous places — among talus slopes, boul- 
der fields, etc. In deserts and semideserts, the weasel is closely 
associated with the places where colonies of social rodents are 
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 local- 
ity. 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 kills its prey. It can bite 
through [the skull of] a young water vole only with difficulty; it 
can deal with pikas and gerbils, but it cannot overcome young 
Norway rats and sousliks. 

Species composition of prey is also determined by habitat of 
the weasel — in the same locality, it behaves different in forest than 
near villages and in the fields. A characteristic feature of the weasel 
is the fact that in its food, shrews are more numerous than in other 
carnivores (Formozov, 1948); in different years in Tatariya, their 
frequency reaches 25% (Grigor'ev and Teplov, 1939). 

In Murmansk district, the diet of the weasel consists of gray 
[Microtus] and red-backed [Clethrionomys] voles, Norway 
[Lemmus] and wood [Myopus] lemmings, and rarely fish 



*Strip of uncultivated land between two fields — Sci. Ed. 

**Two Russian words, tok and gumno, are both translated as "threshing-floor" — 
Sci. Ed. 



988 

(Nasimovich, 1948). In Trans-Baikal, the weasel feeds on narrow- 
skulled 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 it as well 
as 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 west- 
ern Siberia, small voles, field mice, Dzhungarian hamsters, 
659 [Phodopus sungorus], water voles enter the weasel diet, and in 
small amounts, water voles and fish, and even carrion is some- 
times 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 cap- 
tivity, water shrews [Neomys] are very unwillingly utilized (V.V. 
Kucheruk). Frogs, fish, small birds and bird eggs are very rare in 
weasel food. In Ussuri Territory, the weasel often eats terrestrial 
molluscs and feeds on a number of marine organisms cast up on 
the seashore (Yu.A. Salmin and V.O. Shamykin). 

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 ex- 
tremely rarely. In the stomachs of weasels are the remains, at one 
time of only one mouse-like rodent; in it usually occur no more 
than 10 gm 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 the little animal is equal to about 35 gm, i.e. about 30- 
35% of its body weight (V.V. Kucheruk). However, weasel kills 
significantly more mouse-like rodents and shrews than needed for 
food. When its prey is found in abundance, it eats only a small 
part of captured animal, 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 
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 
out from a burrow of a great gerbil, Rhombomys opimus. Kyzyl-Dzhar in 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 among a series of adjacent areas of daily activ- 
ity. The home ranges of males and females for the most part occur 
contiguously. In the forest-steppe of western Siberia, the area of 

660 daily activity sometimes comprises 0.5-0.6 ha (Zverev, 1931). In 
Murmansk district, the 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, 
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 "Тер"., 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. 

There is little reliable data on brood burrows. There are indi- 
cations 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 nest- 
ing 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 
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 in a colony of great gerbil. Kyzyl-Dzhar in 
Badkhyz reserve, southern Turkmeniya. A steppe agarna is 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 an- 
other more prey-rich area. The weasel usually hides for a long 
period in subnival burrows of voles and shrews. Very often, it 
uses the underground passages of moles. In severe frost or crusted 
snow, the weasel almost never appears on the snow surface. It almost 
always catches its prey under the snow and rarely on its surface. 

The weasel moves exclusively by jumps, with the distance 
between tracks of the fore and hind feet from 18-35 cm, depend- 
661 ing on the leap and the size of the animal. In Middle Russia, length 
of the leap is 18-20 cm, the feet stand close in the leap. Given the 
possibility of killing 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 appear- 
ances, all stages in the reproductive cycle of the weasel, except 
possibly the duration of pregnancy (from the moment of placentation 
to parturition) are inconstant and variable. They are determined by 
factors of the external environment, and first of all by food abun- 
dance. 

In Silesia, pregnant females were found in January, April, 
August, October and November (Pohle, 1909); in the Caucasus — 
in April, May and October (Satunin, 1915). In southeastern Trans- 
Baikal, 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 
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. 

Most often, litters of 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 
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 is 19 (V.V. Kucheruk). 

Growth, development, and molt. Weasels are born blind, help- 
less, covered 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. 

Enemies, diseases, parasites, mortality, and competitors. All 
carnivorous animals and birds feeding on mouse-like rodents, and 
first of all the ermine, are competitors of the weasel. Many of 
these are, at the same time, enemies of the weasel, destroying it. 
Remains of eaten weasels occur in excrement and debris of red 
fox, sable, steppe and forest polecats, ermine, eagle-owl, buzzard 
and others. Such cases are rare, but nevertheless occur in places. 

Diseases of weasels are almost unstudied. There is some infor- 
mation that the weasel is susceptible to rodent plague; a culture of 
tularemia infection was isolated from it, but the weasel is quite 
resistant to tularemia (Olsuf'ev and Dunaeva, 1951). Skryabingilosis 
is widespread among weasels, as also in 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. 

Population dynamics. Fluctuations in numbers of weasels is 
well-demonstrated. They are very closely connected with the abun- 
dance of food — mouse-like rodents — not of only one species, but 
662 of the 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 ro- 
dents. 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 number 
of weasels rose three times (P.B. Yurgenson). In Kostromsk dis- 
trict, 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 in numbers is explained by high fertility, and sharp 
fluctuations — by narrow feeding specialization (voles and shrews). 
In southeastern Trans-Baikal, within a nine-year period the number 
of weasels changed by 2-3-fold (P.B. Yurgenson). In the Mongo- 
lian Republic, between 1944-45, there was noted a nine-fold in- 
crease in number of weasels (Kucheruk, 1948). In May 1944, in 
one hectare of the test areas, an average of 0.14 weasel was ob- 
tained; in June — 0.66; in September — 1.27 weasels. In April 1945, 
no weasels were caught in any of the test areas. In Kostromsk 
district, during one year the weasel population increased and de- 
creased 10-fold (Formozov, 1948). During the period from 1930 to 
1950, the weasel population in the southwestern part of Kalinin 
district changed 34-fold (1934/35 and 1947/48; P.B. Yurgenson), 
and in the year 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 
(Teplov, 1951). 

Field characteristics. It is very easy to confuse the track of a 
large male weasel with those of a small female ermine. In such 
cases, the weasel is distinguished by its considerably smaller area 
of daily activity, the shorter daily length of the trail 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 its feet closer to each 
other than ermine, and in short jumps the tracks of "landing"and 
"bounding up" often join in one chain. The length of leaps of 
weasels in the eastern Altai (Dul'keit, 1956) is 15-25 cm, the 



994 




Fig. 240. Paired foot impressions of male Middle Russian weasel, Mustela (Mustela) 
nivalis L., 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) 



Ф 



Fig. 241. A group of tracks of the Middle Russian weasel, jumping easily on four 

legs on the 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 (in ermine, 1.8-2.5). 
The weasel sinks into the snow for about 1.5-2.5 cm. The weight 
load on 1 cm^ of the surface of the track is 7-8 gm (P.Yu.). 

Practical Significance 

The significance of the weasel in the destruction of harmful mouse- 
like rodents is very great. This is, in particular, strengthened by the 
fact that, in years of abundance, it kills tens 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 also. 

In the fur trade, the role of weasel is negligible. It 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 must be pro- 
tected and preserved throughout the year, and its hunting must be 
prohibited everywhere. Its trade should be everywhere prohibited, 
and the accidentally captured weasel must not be used in fur prepa- 
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. Mag. Nat. Hist., 
15, p. 452. Kara-Karyk mountains, Fergana. (?V.G.). 

1896. Putorius arcticus. Merriam. North Amer. Fauna, 11, p. 15. 
Point Barrow, Alaska. 

1912. Mustela lymani. Hollister. Smiths. Misc. Coll., 60, 14, p. 5. 
Tapucha*, on the Chuisk Trail, southern Altai. 

1913. Mustela nippon. Cabrera. Biol. Soc. Espan., 13, p. 392. 
Sinano, on Hondo (Honshu) Island, Japan. 

1914. Mustela kanei. G. Allen. Proc. New Engl. Zool. Club, 5, p. 
58. Nizhne-Kolymsk. 

1922. Putorius erminea var. kamstschatica. Dybowski. Arch. Tow. 
Nauk. Lwow, 1, p. 349. Nom. nudum. 

1922. Putorius erminea var. sibirica. Dybowski. Ibidem, p. 349. 
Nom. nudum. Nee Pallas, 1773. 

1923. Arctogale erminea tobolica. Ognev. Biologich. izvestiya, I, 
p. 112. Тага, Former Tobol'sk governance. 

1928. Mustela erminea transbaikalica. Ognev. Memuary zool. otd. 
Obshch. lyubit. estestvozn., antrop. i etnogr., 2. Sosnovka, 
Barguzinsk Preserve. 

1928. Mustela erminea orientalis. Ognev. Ibidem, p. 15. 
Pokhodskoe S. [village] on the Kolyma [river], 69°04' 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 
praeoccupatum — Kolonokus alpinus birulai Ogn., 1928. 

1932. Mustela erminea ognevi. Jurgenson. Zool. Anz., 98, p. 11. 
Mouth of Taz river, western Siberia. 

1935. Mustela erminea schnitnikovi. Ognev. Zveri SSSR i 
prilezhashchikh stran, 3, p. 37. 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, 1, p. 124. Voevoli Lake, source of Khatanga, 
66** 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. 

1944. Mustela erminea digna Hall. Proc. Calif. Acad. S,c., 23, 
No. 37, p. 559. Kamchatka. 

1951. Mustela erminea martinoi. EUerman et Morrison-Scott. 
Checklist Palaearct. Indian Mamm., p. 256. Substitute for 
M. e. birulai Martino. 

1962. Mustela erminea balkarica. Baziev. Zool. Zhurn., 41, No. 1, 
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 tail black. Tail length with terminal hairs comprises about 
half to more than a third of body length. Skull moderately elon- 
gated — distance between the mastoid processes approximately equal 
to half of condylobasal length, but somewhat more elongated than 
in weasel. Width of skull above canines noteably less than inter- 
orbital space. Dimensions are small (V.H.). 

Description 

In its general proportions, manner of posture and movement, 
the ermine is entirely similar to the weasel and represents a some- 
what enlarged copy of it. However, the tail is relatively longer, 
never being equal to the length of the hind foot, always exceeding 
a third of body length, and its length together with terminal hairs 
constitutes about half of body length. 

Winter fur very dense and silky, but quite closely-lying and 
short; summer fur rougher, shorter and sparse. Among ermine 
around Moscow, length of directional hairs in winter is up to 
17 mm (M15.75), and in summer, M13.9; guard hairs of category 
I in winter to 15 mm (M13.7), and in summer, M12.1; guard hairs 
in category II in winter to 14 mm (M12.2), in summer Ml 1.3; 
guard hairs in category III in winter to 13 mm (Ml 1.9), in summer 
M10.5; guard hairs in category IV to 12 mm (Ml 1.5), in summer 
M9.9; length of underfur in winter M9.1 mm, in summer 8.1. The 
total number of hairs in 0.25 cm^ in winter comprises 4808, of 



998 



665 



which two* are directional hairs while underfur hairs number 
4 637 — 1 for every 27 underhairs; in summer, on 0.25 cm^ there 
are 3,636 hairs, of which there are 5 directional hairs and under- 
hairs number 3,539 — 1 for every 36.5 underhairs. Thickness of 
fur 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 closely- 
lying hairs, not fluffy, and relatively fine and even throughout 
whole length of the tail. 

Soles of feet furred and in winter fur, their pads are not vis- 
ible. In summer, pelage is less dense and pads are bared. 

In summer color is 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 is quite 
bright yellow. The distal half of the tail is black in winter and in 




Fig. 242. Ermine, Mustela (Mustela) erminea L., in summer fur. Sketch by 

A.N. Komarov. 

*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 to quite light straw-brown. Geo- 
graphic variation in color of the winter fur is not expressed. As 
noted, seasonal variation is great and expressed in all races. Only 
a few of the southernmost populations or individual animals in 
these populations do not turn white or partially white in winter. 

The ermine skull is very similar to that of weasel, but 
relatively more elongated, with a less broad and swollen braincase. 
Its anterior region (in the region of the frontal bones) is not so 
wide and voluminous and is more elongated. The postorbital con- 
striction is sharper and somewhat longer. The facial portion of the 
skull is elongated and relatively narrow — skull width above the 
canines is notably less than width of the interorbital expanse. The 
zygomatic arches are weak and thin. Zygomatic width approxi- 
mately corresponds with skull width in the mastoid region. The 
infraorbital foramina are rounded and relatively large — their 
transverse diameter is considerably larger than the longitudinal 
diameter of the canine alveolus. 

The upper profile of the skull in the region posterior to the 
supraorbital processes is flattened, and therefore in the interorbital 
667 region, there is an evident elevation, from which the profile of the 
facial part quite abruptly descends. The prominences, crests, etc. 
are weakly defined, but relatively somewhat stronger than in wea- 
sel — the better defined is the sagittal crest found in the postorbital 
(frontal) region, and the occipital crest is well defined. The audi- 
tory 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 
somewhat stronger than in weasel. Carnassial teeth are well devel- 
oped, the upper canines longer and thinner and the lower canines 
somewhat massive. 

On the whole, because of its relative elongation, less swollen 
braincase and its flatness, elongation of facial part, development of 
crest, prominences, etc., 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 in the ermine skull are quite considerable. '' The 
young ermine skull (first winter of life), besides somewhat smaller 
general dimensions, is 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 
a skull which cannot be differentiated from that of adults in its main 
features (for age changes, see also description of the os penis). 

Sexual differences are considerable and are manifested in its 
somewhat different proportions of the female skull besides its 
generally smaller dimensions. In females, the skull is somewhat 
narrower in the nasal part (about 23.3% of condylobasal length 
against 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- 
als, 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 
the male skull is 165% of that of the female skull. 

Condylobasal length of the male skull (the same series) is 45.5- 
M48. 39-5 1.8 mm, that of females is 40.3-M43.39-45.8 mm (V. 
Popov, 1947). 

Extent of individual variation in skulls is quite considerable in 
general, particularly in characters of general size; however, this 
variability is of "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 
particularly in the postorbital constriction which may be longer 
and wider or shorter, sharper or weaker. All of this leads to the 

'^Age and sexual variation in the ermine skull was the subject of several special 
investigations; it was better studied than other species of the family. It serves, to a 
certain degree, as a model for the genus. Very general information is given here. For 
details, see 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 
and, in general, obviously differ in their general appearance. This 
has repeatedly led to misunderstandings in description of separate 
forms. Geographic variation of the ermine skull is insignificant, much 
less than in weasel, and these differences are never so great. Geo- 
graphical changes are not of the degree seen in sexual dimorphism. 
668 The OS penis differs 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-M25.5- 
27.7 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 0.016- 
MO.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 
foot of males is 40.0-48.2 mm, of females, 37.0^7.6 mm; height of 
ear in males is 18.0-23.2 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. 

Weight of males to 258 gm, of females to 180 gm, usually 
much less'» (V.H.). 



"According to Stroganov (1962) data for ermine of our country. They also char- 
acterize ermine of West Europe. Here, only individual animals (judging from a large 
series; Reichstein, 1957) with perhaps slightly larger measurements (interorbital width 
of males 13.7, females 11.8 mm; zygomatic width of females 24.6). The given ampli- 
tude and dimensions also applies to the small form of ermine from the highlands of 
the western and central Alps (M. e. minima). Some measurements given by Novikov 
(1956) are not fully accurate (body length 160-380 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). 



1003 

Systematic Position 

The relationship of ermine to affiliated species of the genus was 
reviewed in the description of weasel (see above) and M. altaica 
(see below), with which this species is closely related. The most 
fundamental difference lies in the structure of the os penis which 
is apparently, important in a biological sense. In contrast to wea- 
sel, the ermine must be considered as a more "specialized" or 
rather "advanced" form, although the weakening of "infantile" 
features characteristic of weasel, are probably connected primarily 
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 the range constitutes the coast of the 
Arctic Ocean, to which the ermine reaches even in the northern 
Taimyr, i.e. 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- 
ably Beloi, Nordenskjold; in the Laptev Sea — 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 
670 remaining islands of the Arctic (occurrence on Baigach is entirely 
possible). In the east, the border of the range passes along the 
Bering, Okhotsk and Japanese seacoasts and inhabits Karagin 
Island, 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. 
Voronov). It is encountered on Sakhalin and all of the Shantar 
Islands. 

The western frontier, 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 Kazakhstan extends into the steppe and semidesert, 
and in part even the desert zone. In the west, between the state 
frontier and the mouth of the Dnepr, the ermine is met with up to 
the Black Sea coast and is found even on Tendrov Spit. The er- 
mine is absent on the Crimean Peninsula (information of 
Kalenichenko, 1839, is erroneous) and eastwards from the Dnepr 
in the narrow steppe belt, which is directly adjacent to the Black 
and Azov seas. It exists immediately around Askaniya Nova and at 
the mouth of the Don, and perhaps, even below Taganrog and 
along the northern shore of Taganrog Gulf of the Azov Sea. Far- 
ther 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 pre- 
cise data on the southern limit of the ermine's range in the steppes 
of the Cis-Caucasus and at the 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 pre- 
serve'^. 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 to the northwestern shore of the 
Aral Sea, continues along its 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. 
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 this finding did not receive 
confirmation, the occurrence of ermine in the Caucasus was rejected or at least not 
confirmed. The fact that 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. 



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 bound- 
ary 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. 

The range in Middle Asia may be outlined in the following 
form. In the desert lowlands and sands south of [Lake] Balkhash 
and along the rivers flow into the lake, the ermine is absent. It 
exists in the 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 
Alatau. Ermine is absent in the Karatau. 
671 The species described is found in Fergana and in the Pamiro- 

Alaisk system. Details about its distribution here are almost ab- 
sent. 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, 
the ermine 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 coastal zone northwards to a region somewhat north of the 

^"Range according to data of Kalenichenko, 1839; A.M. Nikol'skii, 1891; 
Byalynitskii-Biruli, 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, 
1948, 1948a, 1952; Shereshevskii and Petryaev, 1949; Heptner, Turova and Tsalkin, 
1950; Bazhanov, 1951, 1952; Korneev, 1952; Sludskii, 1953; Bannikov, 1954, 
Podkovyrkin, 1958; Afanas'ev, 1960; Sokur, 1960; Tembotov, 1960; Baziev, 1961; 
Stroganov, 1962 and other sources. 



1007 



mouth of the Samarga. In this region, the range of the ermine 
extends along the main axis of the Sikhote-Alin' southwards ap- 
proximately to 44°30' (the latitude of the southern extremity of 
[Lake] Khanka). In the western half of the territory from the Amur 
to Sikhote-Alin', the boundary of the range apparently passes to 
the middle Bikin in the form of an arched line from the Khabarovsk 
region^'. 

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). It is 
absent in Bulgaria, Albania, Greece and Turkey. To the west it 
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 
northeast of Peshawar (Hazara — the southernmost place of occur- 
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. Undoubt- 
edly, it lives in Great Khingan and, probably, in the northern 
montane regions (Lesser Khingan, Il'Khuri-Alin'), and perhaps also 
in the east. Possibly it is found in North Korea. Its distribution in 
Japan includes at least Hokkaido and Honshu (Hondo). 

The reference concerning its occurrence in Algeria (whence a 
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, 
the ermine does not extend south of the Pyrenees and the Alps, i.e. 
that this form is isolated in its distribution. It is entirely probable 
that this name applies to the large southern weasel (M. п.* 
numidica), as it is considered by some authors (G. Allen, 1939). 
In North America, the range occupies the entire Arctic 
archipelago, including Ellsmere Land (Grant), northern Greenland 
(northernmost point inhabited by the species) and eastern Green- 
land 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 
673 southernmost point inhabited by the species is about 34° N. lat.). 
Thence, the border sharply rises to southwestern Alberta and south- 
ern 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 is- 
lands in the Bering Sea which are located near the Alaskan coast 
(no indications for the Pribilovs) (V.H.). 

Geographic Variation 

In vast area of its range, the ermine reveals some geographic vari- 
ation which, however, is not great. Its amplitude, in a morphologi- 
cal sense does not attain the dimension observed in the weasel, and 
is not more than that in other comparatively slightly variable spe- 
cies of the family and genus. It mainly bears a clinal character. 
The winter coat (its density, length), the color of the summer fur, 
and in part the general dimensions, are chiefly those that vary 
geographically. 

Geographic variation of the ermine is still insufficiently stud- 
ied, and representative of this, it is, apparently, exaggerated. Within 
the last decade, in reviews of our fauna, 10 subspecies were at first 

*In Russian original, erroneously as e — Sci. Ed. 



1010 

accepted (Ognev, 1935); later, although with reservations — 13 
(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 less. 
Several foreign investigators accept many forms — 21 for the Old 
World (Ellermann and Morrison-Scott, 1951) and 20 for America 
(Hall, 1945; Hall and Kelson, 1959), where the extent of the range 
is much smaller than in Eurasia. 

The following forms may be provisionally accepted for our 
country (characteristics mainly after Morozova-Turova, 1961 and 
Stroganov, 1962, with modification). 

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 USSR — Scandinavian Peninsula. 

Kola ermine are poorly known and the actual relationship of 
the 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 vari- 
ation in color is insignificant — from light-reddish to dark-tawny 
tones. 

Body length of males (57), 215-M248.7-278 mm, of females 
(23) is 195-283 mm; tail length of males,78-M92.4-105 mm, of 
females, 60-86 mm; length of hind feet of males 41.5-M46.2-55 
mm. 

Condylobasal length of skull of males (76), 41.0-M46.8-52 
mm, 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 skull, 19.0-M22.9-26.2 mm, 
of females, 18.0-M21.1-24.5 mm Zygomatic width constitutes 
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 (syn. 
balcarica). 

Dimensions small. 

Color coffee- or reddish-tawny. 

*Only mean value in Russian original — Sci. Ed. 



1011 

674 Body length of males 217 mm, of females, 175-190 mm; tail 
length of males, 84 mm, of females, 57-61 mm; length of hind 
foot of males 40 mm, of females, 27-30 mm. 

Condylobasal length of male skull, 43.0 mm, of females, 
37.8-39.3 mm; zygomatic width of males, 22.6 mm, of females, 20.0- 
20.6 mm; interorbital width of males 9.8 mm; of females, 18.4-18.6 
mm. (Baziev, 1962). 

Northern slope of middle part of Main Caucasus range (El'brus 
massif). 

Outside the USSR — absent. 

A very little known form, described as similar in dimensions 
and color to M. (M.) e. 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 large, on average somewhat larger than in the form 
aestiva. 

Winter fur tall, dense and silky. Summer fur is also somewhat 
longer, denser and softer. Skull is large. Zygomatic width consti- 
tutes 54.6-M57.7-58.1% of the condylobasal length of its skull. 

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, 24.8- 
M27.7-30.0 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-M10.8- 
12.0 mm, postorbital width of males, 9. 6-Ml 1.2-12.1 mm, of 
females, 8.2-M9.2-11.1 mm; mastoid width of males, 21.2-M23.6- 
25.2 mm, of females, 1 9. 3-M2 1.0-22.2 mm. 

In western Siberia, eastwards to the Yenisei and Altai, and in 
Kazakhstan. 

Absent outside the USSR. 

^^The minimum of this measurement (Stroganov, 1962) strongly differs from that 
given previously according to the same author (p. 668) and, apparently, here the 
mistake is self-inflicted*. 

*In the Russian original, the word "Kroetsya" is used, which cannot be found. 
It may be a misspelling of "kropatsya", the reflexive form of the verb to bungle — Sci. 
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- 
tures (Ognev, 1935; Morozova-Turova, 1961). According to 
Stroganov (1962), only characteristics of fur distinguish the Tobalsk 
ermine from the form aestiva. 

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 dis- 
appears completely. Skull similar to that of M. e. aestiva. Distance 
between zygomatic arches relatively narrow (zygomatic width con- 
stitutes 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 
females, 40-M60-90 mm; length of hind foot of males, 37-M44- 
48 mm, of females, 32-M36-42 mm. 

Condylobasal length of male skull (14), 44.0-M48.4-49.4 mm, 
of females, 41.2-M43.7-45.2 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-M11.3- 
12.0 mm; postorbital width of males, 10. 0-Ml 1.4-12.8 mm, of 
females; 9.0-M10.2-1 1.7 mm; mastoid width of males, 21.3- 
M23.6-25.3 mm, of females, 19.2-M21. 1-23.0 mm. 
675 Weight of males (5), 145-M191-247 gm, of females (4), 106- 
149 gm (Altai preserve, winter; material from ZMMU). 

Mountains of southern Siberia eastwards to Baikal (Altai, 
Sayan). 

Outside the USSR — in contiguous parts of the Mongolian 
Republic. In the Gobi Altai and southern parts of Khangai and the 
Mongolian Altai, apparently, is another form. 

6. East Siberian ermine, M. (M.) e. kaneii Baird, 1857 (syn. 
orientalis, naumovi, sibirica, kamtschatica, kanei, digna; some- 
times 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 
tinges of various intensities. Zygomatic width constitutes about 
56% on average of condylobasal length of skull. 

Body length of males (22), 213-M260-325 mm, of females 
(7), 176-M212-222 mm; tail length of males, 70-M86-100 mm, 



1013 



of females, 67-M72-77 mm; length of hind foot of males, 40- 
M43— 48 mm; of females, 33-M38-43 mm; height of ear in males, 
20-M21-22 mm, of females, 18-M 19-20 mm. 

Condylobasal length of male skull (24), 44.0-M46.0-49.2 mm, 
of females (23), 3 8. 9-M40. 9-44.3 mm; zygomatic width of males, 
24.0-M26.2-28.8 mm, of females, 20.0-M22.3-25.2 mm; inter- 
orbital width of males, 10. 1-Ml 1.4-13.2 mm, of females, 8.8- 
M9. 5-10.0 mm; postorbital width of males, 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, ex- 
cept 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 somewhat smaller 
dimensions. Differences can only be noted in series. The form is 
close to M. e. arctica from Alaska and may be identical with it. 

7. Karagin ermine, M. (M.) e. 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, 9.8- 
MlO.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 con- 
firmed 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 fur short and sparse, dark-brown in color. Skull small 
with relatively narrow interzygomatic area — zygomatic width con- 
stitutes on average 51.6% of condylobasal length of skull. 

Body length of males (14), 225-242 mm, tail length 80-M85.7- 
90 mm, length of hind foot, 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; inter- 
orbital width of males, 9.1-M10.8-11.6 mm, of females, 7.9-M8.7- 
10.0 mm. 
676 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 
mm, 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] 
U[niversity]). 

In Trans-Baikaliya, Amur area, Ussuri Territory, Shantar Is- 
lands. 

Outside the USSR probably in the eastern part of the Mongo- 
lian Republic and northeastern China (former Manchuria). 

This form is close to the Altai, M. (M.) e. lymani and possibly 
both may comprise one form, which occupies all of southern 
montane Siberia and above-mentioned parts of the Far East. The 
name lymani has priority. 

9. Fergana ermine, M. (M.) e. ferghanae Thomas, 1895 (syn. 
schnitnikovi) . 

Dimensions small. 

Color of summer fur very light, straw-brownish or grayish, 
rarely with a film of red. The coat is short and soft. On the neck 
occur light spots, sometimes forming a collar. This is the lightest 
colored form of our fauna. Individual animals do not turn white in 
winter or become only partially lighter. 

Body length of males, 205-M259-300 mm; tail length 70- 
M75-82 mm; length of hind foot, 40-M41-42 mm. 

Condylobasal length of male skulls (9), 40-M43-45 mm; 
zygomatic width, 22.0-M23.3-25.5 mm; interorbital width, 9.5- 
M 10.4-1 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 India 
and the westernmost parts of Tibet; found in adjacent parts of Tien 
Shan China. 



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 parts of 
the range lying outside the boundaries of the USSR, is great and 
evidently exaggerated, particularly for America. 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- 
graphic distribution"); 3) M. (M.) e. stabilis Barrett-Hamilton, 
1904 — England; 4) M. (M.) e. ricinae Miller, 1907 — islands of the 
Hebrides; 5) M. (M.) e. minima Cavazza, 1912 — Switzerland; 6) 
M. (M.) e. nippon Cabrera, 1913 — Japan; 7) M. (M.) e. mongolica 
Ognev, 1928 — Gobi Altai (apparently, a synonym of ferghanae); 
8) M. (M.) e. arctica Merriam, 1896 — Alaska, northwestern parts 
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 main- 
land; 11) M. (Л/.) e. richardsonii Bonaparte, 1838 — Newfound- 
land, 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 
of the species in America (Nevada, Utah, Colorado and other states); 
15) M. (M.) e. kadiacensis Merriam, 1896 — Kodiak Island; 16) M. 
677 (д^.) e, 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 Alexan- 
der 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 the number of abundant car- 
nivores. Within the boundaries of its range, its numbers and den- 
sity are, however, distributed very unevenly. 

Indices of prepared skins, even those calculated per unit area 
(usually per 10 km^) by no means reflect the actual condition of 
the species population. 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 popu- 
lation 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. It is more frequent in the steppes of 
southern Ukrainian SSR and in the southeastern European part of 
the USSR (P.B. Yurgenson). Here, it is 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, Siberia— 1.07, Tatar 
ASSR— 0.78, Ural— 0.85, Severnyi Territory— 0.60. In other parts 
of the range, abundance of ermine is considerably lower. 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 main regions where the ermine was abun- 
dant, its numbers 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 'of rivers, 
riparian meadows and thickets of bushes. It also lives along the 
slopes of valleys and in rocky places. The latter are particularly 
typical for montane tundra. It also willingly holds to the fringe of 
the forest-tundra. In the forest zone, 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 
thickets and hummocks, etc. Forest massifs are untypical habitat 
for the ermine, although here it is met with almost everywhere, but 
not often. In forests, old cluttered burned areas and glades, the 
678 edges of forests and shrubby areas (especially those near settle- 
ments and arable lands), but deep within forest massifs it prefers 
cultivated fir and alder as well as the forest meadows and old hay- 
stacks. It does not avoid settlements and gardens near them. In win- 
ter, 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 in the forest and to 
decrease in the area inhabited by it (for instance, 100 times). As 
a result of this process, the greatest density of ermine occupation 
in fir forests is found in the cultivated fir plantations which are 
teeming with life. These are original surviving stands, whence under 
favorable conditions ermine spread out into other stands. Flooded 
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 clear- 
ings and their edges, in birch-groves along swamps, in hummocky 
swamps and in weedy thickets around steppe lakes. It is also 
encountered along steppe ravines and gorges. It often appears in 
dwellings not only of small settlements, but also the outskirts of 
cities. 

In the Altai mountains, ermine occupies rock slides with pika 
colonies, and dry alpine meadows inhabited by the narrow-skulled 
vole [Microtus gregalis]. Here, it more willingly lives in places 
with shallow snow cover. In selecting stands, the ermine is more 
plastic than sable and Siberian weasel. It is more rarely met with 
in dark coniferous taiga than in thin, sunlit stands. On coastal 



1018 

cliffs it lives in rock slides, 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 
to stands with willow ptarmigan and lives for a long time in rock 
slides inhabited by pikas (Dul'keit, 1956). In the mountains of 
southern and southeastern Kazakhstan places of its occupation 
include shrubby thickets, apple, juniper and fir forests and high- 
land fescue steppes (Sludskii, 1953). The ermine everywhere pre- 
fers floodlands to other types and without them, it spends much of 
its time near water. In Tatariya, in flooded areas, the density of 
ermine is six times higher than in plakorn* dry regions (Aspisov 
and Popov, 1940). 

Food. According to feeding type, the ermine belongs among 
animals with a limited assortment of food. In its food mouse-like 
rodents (in the broad sense) predominate, but in contrast to the 
weasel which almost exclusively feeds on small voles, a consider- 
able place in its food is occupied by the larger rodent species — 
water vole, common hamster, pikas and others, which the ermine 
overpowers in their burrows and which the weasel cannot master 
or with great difficulty. 

More or less constant, but of secondary significance, are small 
birds, and sometimes fish and shrews. More rarely there are am- 
phibians, 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 
destroyed by ermine were recorded during four years. Of these in 
60 cases it was nests of the pied flycatcher, in 59 — starlings, in 
16 — great tit, in 8 — spotted flycatcher, in 6 — wryneck, and in 4 — 
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 

the water vole and gray vole [Clethrionomys rufocanus], the root, 

common and field voles [Microtus oeconomus, M. arvalis, M. 

680 agrestis] predominate; in forest areas, various species of red-backed 

voles; in tundras, lemmings also. Ermine rarely attack the commercial 

*Loca{ word? — Sci. Ed. 



1019 



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1020 

animals and birds — even more rarely with success. However, cases 
of ermine feeding on muskxats, probably young, have been recorded. 

Geographic variation in nutrition of ermine is not great. Vari- 
ations 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 
eats no more than two and in a famine year, one satisfies it. With 
an abundance of prey, it consumes only the brain or leaves it 
untouched on the trail, killing significantly 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 
hamsters, 5-8 water voles, up to 10 small mouse-like rodents and 
4 shrews at one time, etc. (Zverev, 1931 and others). 

Home range. With the formation of snow cover deeper than 
12-15 cm 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 




Ri 


inge of fluctuation 
in occurrence 


Average for 
1938/39 to 1950/51 


Voles 






0—72.7 




47.9 


Shrews 






5.9—66.9 




24.2 


Tetraonid birds 






0—13.3 




7.8 


Other birds 






0—35.8 




11.8 


Squirrel 






0—11.2 




1.9 


White hare 






0—7.4 




0.7 


Carrion 






0—6.7 




1.5 


Insects 






0—7.1 




0.5 


Percentage of empty 


stomachs 


29—58 




46.8 


Average weight 


of stomach 


0.5— 3.0g 




1.35g 


contents 













Note. From data of the "Nature Chronicle" of Pechora-Ilych preserve. 



1021 

This area is divided into a series of ranges of daily activity, 
with repeated visits each 3-10 days (Zharkov, 1941). Such ranges 
of mingled living animals often overlap each other. 

In sparsely treed, northern montane Murmansk district, the area 
of the home range varies within limits of 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 it reaches 100-200 hectares 
(Shibanov, 1935). In the floodlands of Tatariya, the area of the 
daily activity range varies 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 
of ranges are observed, inhabited by 5-12 individuals and alternat- 
ing with ranges of lower density of occupation. It is possible that 
these are mingled ranges of the dispersing litter (Zharkov, 1941). 

Burrows and shelters. The ermine does not dig a burrow inde- 
pendently. It often uses the burrows and nest chambers of the 
rodents it kills — water voles, hamsters, Siberian chipmunks, and 
others — for the litter young, 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 unex- 
pected and seemingly unsuitable places, for example among logs 
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 rest. Such shelters occur under rocks, in the roots 

*There is an obvious misprint here, but what the correct values are could not be 
determined — Sci. Ed. 



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 is mostly active in the 
twilight and night hours of the diel period, but sometimes it comes 
to pass that it is encountered during the day. The light regime in 
high latitudes substantially influences the diel cycle of the ermine. 
In the period of the polar day, about 70% of the occurrences of 
this animal take place in the night hours (1800-2400). In autumn 
(September-October), when the night is becoming darker, it often 
is 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 — 11 encounters, day — 7, evening and 
night — 5; Nasimovich, 1948). 

During periods of severe frosts activity of the ermine notably 
decreases. With its 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, 1931; Zharkov, 1941; 
Nasimovich, 1948). In winter, it goes out for prey 1-2 times per 
day. 

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, 
1948). 

The ermine swims and climbs well, but it is fundamentally a 
terrestrial animal. It can move freely in the underground passages 
of rodents — water voles, hamsters and pikas, but it is not able to 
penetrate the burrows of small voles. The ermine is very bold, 
brave and cunning. The prey is killed 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 
peoples of the far north it often makes itself an intrusive lodger, 
plundering stock of meat, fish and other products. 

The movement of the ermine is quick and dexterous, but some- 
what 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 is typical for ermine in winter in years when 
it is sufficiently supplied with food; when unbalanced by 



1023 

starvation or insufficient food, then it is obliged to roam widely. 
In Lapland preserve, an ermine roamed an area of 25 km^ 
(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 com- 
pared with mass reproduction of water voles in the northern re- 
gions (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 — grain 
stores, straw piles 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- 
tions from the pine woods and burns to the fir groves for a dis- 
tance of 10 to 15 km were observed (Nasimovich, 1948). 

Reproduction. Reproduction of the ermine is still entirely in- 
sufficiently studied. It has one sexual cycle per year. Spermatogen- 
esis in males lasts for five months of the year, and sexual activity 
for four months (middle of February-first half of June for northern 
Kazakhstan). In the nonfertilized females, ovulation is periodically 
repeated monthly. They are capable of being fertilized throughout 
the whole period of male sexual activity. In adult females, 
fertilization for the most part takes place just after parturition. 
Young females are fertilized in summer. During the course of 
pregnancy, there is a prolonged latent period, probably with 

* Local name? — Sci. Ed. 

** Misspelled Severtsev in Russian original? Not in Lit. Cit. — Sci. Ed. 



1024 

variable duration. After it, the active development of the embryos 
lasts not more than one month. 

The number of young in a litter varies from 2-18, more often 
4-8; for northern Kazakhstan, the average number of young is 8.7 
(Tikhvinskii, 1937; Grigor'ev, 1933; Lavrov, 1944). In Yakutiya, 
estrus usually occurs in April-May, but sometimes even in the first 
half of April, embryos are observed easily by eye. Their number 
ranges from 7-8 up to 12. In Yakutiya, the number of males in a 
litter is greater than that of females (Belyk, 1958). In populations 
of West Siberia, males constitute 53% and females — 47% (Zverev, 
1931). In northern Kazakhstan, males are 61-65% of the popula- 
tion; 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 ermine of Yamal, two peaks of reproduction are observed: 
in late 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 in April-May. At that time, those 
females who have not, for various reasons, participated in the sum- 
mer 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 is subject to 
geographic variation, but this question has not been studied. 

Growth, development and molt. The young are born in March- 
April, blind, 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 as well as the Siberian 
weasel occupy successional habitats which are sometimes isolated 
from the habitats of sable, and the latter animal occupies the widely 
distributed primary habitats. The ermine coexists better with sable 
than does the Siberian weasel. Dispersal of ermine takes place at 
the end of November-beginning of December when the sable has 
already determined 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, 
1959). 

Diseases of the ermine are insufficiently studied. The ermine 
is 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 tape- 
worms 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 patho- 
logical significance. In different years, the degree of infestation 
reaches 50% of the population, and the number of individual worms 
in one ermine — up to 45. Beside mortality, skryabingulosis causes 
a reduction in fertility. Tapeworm invasion of the ermine is 
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 37.4% (average 21.3%). In evaluating these 
figures, selectivity of the harvest must be taken into consideration. 
In Tatariya, the quantity of juvenile animals varied over 11 years 
from 19.3 to 63.9%, constituting an average of 50.8% (V. Popov, 
1947). In 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 
winter, young ermine constituted 94.4% of the population, and 
684 adults — 5.6% (2,373 specimens). These data speak to the higher 
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; 
since in autumn, the number of mouse-like rodents — the main 
food of ermine — is highest. 

In Murmansk district, the principal reason for ermine mortality 
is starvation (Nasimovich, 1949). In Kazakhstan, ermines die in 
great numbers during the time of great steppe fires; in connection 
with drying up of water bodies in the steppe; during flooding of 
the steppe from snowmelt water in spring; from hunger as a result 
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 (V.I. Tikhvinskii). 

Harvesting has been shown as a very powerful effect on the 
process of mortality in populations. Its 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. With abundant food, ermine are numerous, but rarely 
enter traps, and on the contrary, they go into traps well and are 
caught in years of starvation with little food. Hence, the increase 
in 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 mouse- 
like rodents may be competitors of the ermine. 

Population dynamics. The ermine population is 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 in conditions of 
the external environment. 

In the process of these changes, numbers of ermine may change 
within different limits: in Murmansk district, up to 30-fold 
(Nasimovich, 1948), in the Tatar ASSR 54-fold (Aspisov and Popov, 
1939), and in the southwestern part of Kalinin district (20 years of 
observation) by more than 189-fold (Yurgenson, 1959). In the first 
8 years of observation, fluctuations here were only 2-6-fold; in the 
post-war years, the population decreased very sharply, as was also 
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 
interval of 3-4 years between two peaks in ermine numbers in 
Murmansk district, 5-6 years in the southwestern part of Kalinin 
district and in the upper Pechora — 6 years etc., there are, appar- 
ently, more prolonged fluctuations within which short-term fluc- 
tuations 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). 
685 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 begin- 
ning of the harvest was 52.0 per 10 km^ , and by its end — only 18.5 
(Zharkov, 1941). 

The reasons for the changes in numbers may be: 

1) changes in availability of its main foods (voles, water voles); 
2) helminth invasion (chiefly in 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 is thus of two kinds: 
moderate floods are favorable, causing a considerable concentra- 
tion of mouse-like rodents in unsubmerged parts in the period when 
young are being fed, but deep flooding is fatal; 4) steppe fires, 
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 snow surface. The ermine moves almost exclusively in jumps, 
leaving on the snow paired, sometimes, somewhat obliquely situ- 
ated, footprints. Tracks and jumps of the males are always larger 
than those of females, but the tracks of young males may be con- 
fused with those of adult females. 





Ф 




Ф 




Ф 



685 Fig. 246. Tracks of large ermine on loose snow lying on a crusted surface. Gait 
unhurried — the prints are situated as a quartet. On the right, the scheme of ordinary 
jumps with paired tracks. Neighborhood of Gorkii. 30 December 1916. Sketch by 
A.N. Formozov, about 2/3 of nat. size. 



1029 

The tracks of ermine differ from those of weasel by their 
larger size, a much larger search route and by the fact that er- 
mines, when walking, sometimes "trots"; i.e. leaves 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- 
40 to 82-100 cm; in weasels their length is 25-36 cm, maximally- 
45 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 
10 cm (P.Yu.). 

Practical Significance 

686 The ermine is one of the fundamental items of mass commercial 
hunting of our country. Its catch in the USSR constitutes not less 
than half of the world catch. Beside this, our country offers ermine 
skins of the highest grade to the world fur market (Berezovskii, 
Ishimskii and other standard fur types). The best grade of the North 
American ermine may only be compared with the 9th grade in 
quality type of our standard. 

Ermine fur is exclusively used for decoration. Because of this, 
its demand and price depend, to a great degree, on style and is 
distinguished by its significant instability. 

Regions where ermine harvest has grown up as a specialty are 
relatively few. It is significantly more often trapped incidentally, 
or near the vicinity of villages. Therefore, the ermine is, on the 
whole, exploited around our country insufficiently and unequally. 

All methods for capturing ermine 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 its fur, which lowers the price and quality; blood- 
staining is also unavoidable when an ermine is caught and throttled 
by a dog. 

Of the passive means of snaring, metal-arch snap-traps, 
dead-fall box traps, kulemka*, and box traps actuated by strong 
elastic — cherkan* — are successfully employed. Less widely dis- 
tributed are various nooses (most often, of hair) which are some- 
times mounted on a frame or crosspiece (called a "little stool"). 
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, 3, p. 98.; 

Altai. 
1823. Putorius alpinus Gebler. Memoires Soc. Imp. Natur. Moscou, 

6, p. 212. Ridder mines, Altai. 
1857. Mustela temon. Hodgson. J. Asiat. Soc. Bengal, 26, p. 207. 

Sikkim, Himalayas. 
1870. Putorius astutus. Milne-Edwards. Nouv. Arch. Mus. Hist. 

Nat. Paris, 7, Bull., p. 72. Mupin in Sichuan, China. 
1911. Mustela longstajfi. Wroughton. J. Bombay Nat. Hist. Soc, 

20, p. 931. Teza, upper Sutlej valley, Himalayas. 
1914. Mustela sacana. Thomas. Ann. Mag. Nat. Hist., 13, p. 566. 

Vic. of Przheval'sk, Tien Shan. 
1928. Kolonocus alpinus raddei. Ognev. Memuary Zool. otdeleniya 

Obshch. lyubit-estestvozn., antrop. i etnogr., 2, p. 9. 

Kulusutaev post near Zun-Torei Lake (Tarei-nor), southeast- 
ern Transbaikaliya. 
1928. Kolonocus alpinus birulai. Ognev. Ibidem, p. 10. Lyangar in 

West Pamir (northern part of Darvazsk range, Obikhangou 

river) (V.H.). 



*Local names of traps — Sci. Ed. 

^'The name is derived from the Mongolian solongo, and is current in Transbaikal. 
In some places, the animal is also known as "suslennik" and "mountain polecat". In 
the fur trade, the name "mountain kolonok" is employed. 



1031 

Diagnosis 

Color from bright ocher-reddish to straw-ocher or from nut-brown- 
ish to brown. Lighter below than above, transition from dorsal 
687 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 
constriction with abrupt isthmus almost immediately behind 
supraorbital processes, its lateral outlines not parallel. Dimensions 
relatively small — condylobasal length of male skull less than 55 
mm, of female, less than 48 mm (V.H.). 

Description 

In general appearance, dimensions and habits, the solongoi is very 
similar to ermine, but differs in its somewhat larger head, longer 
and fluffier fur, the more furry tail. Tail length more than 1/3 body 
length — with terminal hairs, about 1/2. Claws white. 

Winter fur dense, and although short (length of guard hairs on 
sacrum 12-18 mm — only slightly more than ermine), quite luxuri- 
ant. Tail covered by hairs of equal length and equal thickness over 
its whole length as in ermine, but somewhat more luxuriant. In 
some forms, tail hairs very strongly lengthened, even more luxu- 
riant. 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 tips. In middle of back, color more intense. Upper part of 
head is somewhat darker and duller than back (here yellow tone of 
underfur drops out), but "little cap" very weakly defined in form 
of darkish overlay, sometimes not developed at all. 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 
found behind corner of mouth. Tail same color as back, with red- 
dish tint, similarly colored above and below, i.e. darker than ven- 
ter. 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 or whitish. Covered ventrally by dense whit- 
ish-silvery hairs which conceal callosities. 

Summer fur (Altai) considerably shorter, coarser and more 
sparser than winter fur. Hairs on tail considerably shorter and 
sparser, causing it to appear thinner. Lower surface of paws more 
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 
688 somewhat lighter than back — nut-brown. Lower body light- 
ocherous, 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 surface of paws dark, like back, upper has color of 
sides, fur on digits whitish above. Sometimes, small (about 1 cm 
in 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 white or whitish. This color on lower jaw 
imperceptibly passes into color of chin, throat and neck. On upper 
lips, a narrow white field usually quite sharply demarcated from 
dark color of head. Sometimes, white color of upper lips poorly 
developed — narrow or weakly lightened and gradually merges with 
dorsal color. Behind corner of mouth, a bit below border between 
top and bottom, a small chestnut dot occurs. 

Individual variation in color is usually of a fluctuating type 
and of small amplitude. Sexual dimorphism in color is absent. 



1033 

Young up to two months are covered by a short, fluffy, dull, dark 
fur. In the first winter, they are clothed with adult pelage. Geo- 
graphic variation in color is quite significant and is 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 
of the Siberian weasel [M. sibirica], but differs from it in a series 
of essential characters. It 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 sag- 
ittal plane of the skull). 

Infraorbital foramina small, oval, their longitudinal diameter 
less than or equal to longitudinal diameter of upper canine 
alveolus. Interpterygoid area usually narrows forwards, and its 
general outline has an angular form, apex directed forwards. Au- 
ditory bullae are elongated, in plan a smooth rectangular form, 

^■•This distinction between the color of the summer and winter fur is neither 
noted nor emphasized in our 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, its systematic 
position and geographic variation, but also 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 its length. Distance between bullae 
usually greater than width of the hard palate at anterior part of 
690 interpterygoid depression. Zygomatic width approximately corre- 
sponds to width of skull in mastoid region. Zygomatic arches 
weak. Protuberances, crests, etc. unnoticeable; sagittal crest weakly 
defined, occipital crest somewhat larger. Frontal area elevated, 
profile of facial region slopes down from it quite suddenly. Teeth 
relatively strong. 

On the whole, the solongoi skull by comparison with the Si- 
berian weasel skull with which it is usually affiliated, has several 
infantile features. As regards degree of its 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- 
tioned above for ermine. 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, 
but in the second winter of life, the skull acquires all the features 
of the adult animal. Individual variability in the skull is 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 race. 

Structure of the os penis is 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 
length of males 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 
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 23.1- 
27.6 mm, of females, 19.0-24.2 mm; interorbital width of males 
is 8.8-12.2 mm, of females, 7.7-9.5 mm; mastoid width of males 
is 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) M. 
kathiah. It is sometimes considered, and is possibly true, that it is 
only a race of M. altaica. Relationship to the Malayan M. nudipes 
is insufficiently clear, but they are probably close. 

Among our forms and in our literature, M. altaica is accepted, 
according to a tradition proceeding from Satunin, to be closest to 
the Siberian weasel, M. sibirica. This is emphasized by combining 
them into one subgenus (in extreme cases — 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 consid- 
ered (as in keys) to be color. However, there are very essential 
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 
tone. External similarity to the Siberian weasel is found in only 
one race of the species (M. a. raddei) and only in winter pelage. 
691 At the present time, it is more correct to consider that the 

solongoi is, according to several external characteristics and cra- 
niological peculiarities, closer to ermine. It is necessary, however, 
to have in view that as a consequence of the significance of some 
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 be- 
tween 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 more suitable natural relationship 
and refuse to accept the group Kolonocus in any form or rank, or 
in 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 and occupies montane, 
rarely level regions of Central Asia and adjacent parts of Middle 
Asia and Siberia, the Far East and China. 

Geographic Range in the Soviet Union 

In the USSR it is small, and represents the most northeastern, 
northern and northwestern fringe of the species range. It consists 
of several, in part very small, sections separated from each other 
within our country but united to the south, outside the border of 
the USSR. The details and even some of the essential features of 
solongoi distribution are still poorly known and the range cannot 
be outlined with sufficient accuracy. 

In the west, the solongoi inhabits the Pamir, occupying both 
eastern and western parts of the region and the Trans-Alaisk and, 
apparently, the Alaisk range. The northwesternmost known point 
of occurrence in the Pamir is Lyangar in the 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; 
Ishunin, 1961). It is also, apparently, absent in the Turkestan range. 

In the 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 USSR may be isolated from each other. 

In the 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 is remarkable. 

Farther to the east, it lives in the Tarbagatai, Saur, in Zaisan 
depression and the mountains surrounding it, along the Chernyi 
Irtysh and in the Kalbinsk Altai (left 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 its 
occurrence around Barnaul applies, apparently, not to the city it- 
self but to the Altai mountains south of it. 

To the west of the Katun' the solongoi lives, apparently, all 
over the mountains and foothills as far as the Irtysh; it is 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 
(Beskaragaisk region of Pavlodarsk district). In this way, the Altai 
region of occurrence is united with that of Turkestan in our country. 

Deep in the eastern part of the Altai to the east from the Katun', 
the solongoi is, apparently, absent (Turochak lies in the northern 
. foothills). Thus, it 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. 



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1040 

In the literature, very general references to the occurrence of 
the solongoi "in Altai and in Sayan" are occasionally encountered, 
and sometimes, its occurrence in all of these montane regions is 
emphasized. As is apparent, in the Altai the picture is complicated, 
and as regards the western Sayan, not a single accurate positive 
record is found, but there is negative (Stroganov, 1962, after 
Nadeev). There are also no positive data concerning Tuva 
(Yanushevich, 1952, refers to it in this district with a question 
mark; see also the section "Geographic Variation" and notes on 
the Trans-Baikal solongoi). 

The Cis-Baikal part of the range of the solongoi* west of [Lake] 
Baikal is 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 extrem- 
ity 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, 
are true then, it is evident that these — the Eastern Sayan, Cis- 
Baikal and Trans-Baikal areas (see below) of the species distribu- 
tion 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 
beyond the state frontier, except that it 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 

perhaps somewhat beyond it, i.e. to 60** N. lat. and somewhat to 
the north (the northernmost place inhabited 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. 

The distribution to the east is also not accurately established. 
In the northeast, the solongoi is known from the Aldan (Tugarinov, 
Smirnov and Ivanov, 1934), apparently from its upper reaches, and 
in the south it undoubtedly reaches in the east the confluence of 
the Shilka with the Argun' and is recorded in the western parts of 
the Amur district (Plyater-Plokhotskii, 1936)". 

694 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 
not referred to at all and is usually considered to be absent there. 
In actuality, the range apparently covers the whole expanse be- 
tween the Amur and the Stanovoi range, perhaps without reaching 
it, but, occupying the upper reaches (and the sources) of the Zeya 

695 and Selemdzhya. From here, it descends abruptly to the south, 
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^^ somewhere a little to the west of 

693 ^'In the description of the solongoi range east of Baikal, besides the sources 
mentioned, unpublished information offered by 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 
solongoi is very rare everywhere. In the faunistic literature concerning the territory, 
the solongoi is nowhere mentioned. Therefore, it is possible that the range here is in 
fact more restricted than was mentioned. 



1042 



30 60 




694 Fig. 249. Species range of the solongoi, Mustela (Mustela) altaica Pall. (V.G. Heptner). 

132° E. long, or along it, the range border crosses the Amur and 
exits into China. It again enters the borders of the USSR in Primor'e, 
at the Bikin Basin in Ussuri. Thence, the range extends as a 
narrow strip southwards along the Ussuri valley and the adjacent 
western foothills of the Sikhote-Alin', and then, a bit south of the 
latitude of the southern extremity of Lake Khanka, the border turns 
slightly to the southeast and passes to the Ocean at Nakhodka. 

Within the borders of our country, a clear break in the range 
is 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 as well as in the remaining parts of the country, the 
solongoi is absent). In the Korean Peninsula, it inhabits at least its 
northern parts. In China, the range occupies its northeastern part, 
extends eastwards to the middle parts of Shansi and to Hubei (ap- 
parently, to the Great Chinese Plain, from whence it is not re- 
corded), and southwards to Sichuan, Yunnan and Fujian, as well as 
Tibet. In the south, the range includes the Himalayas from Sikkim 
to 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. It is noteworthy that despite the relatively small 
territory occupied in our country by the range of the solongoi, the 
number of these races is quite large, and morphological differ- 
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 some qualitative differences. Due to the scar- 
city of material in our museums, geographic variation of the 

^«Range in the USSR after Maak, 1859; Shrenk, 1859; Radd, 1862; Przheval'skii, 
1870; Gassovskii, 1927; Emel'yanov, 1927; Kuznetsov, 1929, 1948, 1950; Ognev, 
1931; Tugarinov, Smirnov and Ivanov, 1934; Flerov, 1935; Zolotarev, 1936; Plyater- 
Plokhotskii, 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 

solongoi is still, 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]; measure- 
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, 
transition from color of back to color of belly gradual. 
696 In summer 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 
smoky tints of underfur not present). Pure white, or with light 
straw-colored tints on lips, chin and upper part of throat washed 
away at edges, rarely sharply demarcated from color of throat and 
cheeks. Color of lower surface of the body (belly, chest and neck) 
very light, pale-straw or ocherous-white, in some animals almost 
white (bases of hairs often white) with yellowish or reddish over- 
lay, particularly obvious on sides of neck. Dark color of dorsum 
extends to sides very sharply demarcated from light color of ven- 
ter. 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'). 

Body length of males is 224-282 mm, of females, 218-230 
mm; tail length of males is 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 
(lower Hi, Pri-Balkhash; Sludskii, 1953). 

Mountains of Middle Asia including Dzhungarsk Alatau, Pri- 
Balkhash. 

Outside the USSR — probably in contiguous parts of Kashgariya, 
perhaps in Dzhungariya and Afghanistan. 

It is a very sharply distinguished form (material from Tersk 
Altau), the lightest form within the USSR both in summer and in 
winter. In general tone of summer fur and contrast between dor- 
sum and venter of the body it is very reminiscent of ermine. 

Nomenclature of this form is not completely clear. It is un- 
doubtedly (contrary to Ognev, 1931) well differentiated from the 
Altai [form]. The identity of the Tien Shan and Pamir solongoi 



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 is 
similar and there are analogies to such distribution found among 
other mammals. Therefore, the same sacana given to the 
Turkestanian montane solongoi is provisional. 

The systematic position of solongoi of the river valleys in 
Pribalkhash'e is not studied, but its position in this race is 
sufficiently evident. 

2. Altai solongoi M. (M.) a. altaica Pallas, 1811 (syn. alpina). 
Color of winter fur of dorsum darker than in preceding form, 

light ocherous-chestnut color, dull. Lower surface of body much 
lighter than dorsal. Transition 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 
females, 104-M108-113 mm; length of hind foot of males, 41.0- 
M44. 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 (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 USSR — western part of Mongolian Republic (Mon- 
golian Altai). 

A well characterized form, obviously distinguished from both 
eastern and western forms. Darkest form in USSR. 

3. 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 

but also light-ocherous. On sides, transition 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, some- 
what more elongated and sturdy, with narrower interzygomatic 
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, 40.8- 
M42.3-44.2 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; interor- 
bital width of males, 9.3-M10.3-11.8 mm, of females, 8.1-M9.2- 
10.0 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] 
R[epublic], and northeastern China. 

The Trans-Baikal solongoi represents a very well character- 
ized form. In winter pelage, it is the brightest and most beautiful 
solongoi. Summer pelage is a little lighter than in the neighboring 
Altai and 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. Characteris- 
tics of solongoi of the western Sayan are not known, nor is the 
systematic position of solongoi of Ussuri Territory. 

The sharp differences between Altai and Trans-Baikal solongoi 
confirm the opinion about the absence of the species in the eastern 
parts of the Altai 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 M. (M.) a. temon Hodgson, 1857* 
(syn. astutus. longstaffi) is usually recognized, from the Himalayas 
and Tibet. 

Geographic variation in solongoi occurring outside the 
borders of our country demand reexamination, and nomenclature 
revised. Inclusion of "M. kathiah" (with the subspecies caparioccoi) 
in species M. altaica is rejected by the majority of authors (V.H.). 

Biology 

Population. Within the limits of its range, the solongoi is a widely 
distributed and usual, but 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. The most significant part of the range lies in montane 
698 regions where there is negligible development of forest vegetation 
and a predominance of open landscapes. Habitats of solongoi are 
extremely varied. They are, first of all, determined by interrela- 
tionships with the other small carnivores — by 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 in 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). 

In the mountains of Middle Asia, the solongoi is typical of 
altitudes 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 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 
alternate with small lakes, salt flats, ridges and sandhills, the 
solongoi is 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]. 

In western Trans-Baikaliya, the solongoi is common in steppe 
and forest-steppe sections. In upland steppes, the solongoi is char- 
acteristic 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, settling in store- 
houses and dwellings. 

Food. Rodents and birds of moderate and small size, lizards and 
in part, fish constitute the 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; 
rodents consists of 73.2%, birds — 22.7% and insects — 3.1% (97 
spec). Daurian pikas [Ochotona dauricd] occupy first place in 
order of frequency (see Table 65). 

The narrow skulled vole, Brandt's vole and Daurian pika 
predominate 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 Winter Summer* 

27.2 

0.9 

27.2 

36.5 



Daurian pika 


34.2 


Small birds 


13.4 


Brandt's vole [Microtus brandti] 


9.3 


Narrow-skulled vole [M. gregalis] 


9.3 


Daurian partridge[Par^A: dauricae] 


8.2 


Michnoe's vole [M. fortis michnoi] 


6.2 


Clawed jird [Meriones unguiculatus] 


5.1 


Large Asiatic wood mouse [Apodemus 


5.1 


peninsulae] 




Shrews 


3.1 


Daurian hamster [Cricetulus barabensis] 


2.1 


Fish 


1.0 



9.1 



♦Summer food data after material from southeastern Trans-Baikaliya (Daurian 
steppes). 



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 



Summer 



Mammals 


61.0 


Rodents 


61.0 


Voles, mice 


24.2 


Muskrat 


14.7 


Tamarisk jird. [Meriones tamariscinus] 


10.5 


Midday gerbil [M. meridianus] 


1.0 


Water vole [Arvicola terrestris] 


5.8 


House mouse [Mus musculus] 


— 


Tolai hare [Lepus tolai] 


— 


Birds 


5.2 


Pheasant [Phasianus colchicus] 


— 


Moorhen [Gallinula chloropus] 


— 


Ducks 


— 


Small birds 


2.1 


Lizards 


— 


Frogs 


— 


Fish 


3.1 


Insects 


— 


Coagulated blood 


32.6 



66.6 
66.6 

25.2 

13.9 

5.5 

8.7 

11.1 

8.3 

11.1 

2.1 

5.5 

2.7 

2.7 

22.5 

2.7 

8.3 
22.8 



Total data 



95 



36 



Home range. Data are absent. 

Burrows and shelters. In the way of temporary burrows, and 
for rearing young, the solongoi 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 tem- 
porary 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 
activity 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. 



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 to more detailed investigations (Gusev, 1955), the 
first signs of spermatogenesis in individual animals (210 spec.) 
were observed at the end of December. In the first half of January, 
it was observed in all males. Extinguishing of the process 
extended from the second half of March to the first days of May. 
In individual old females, estrus appeared in the first half of Feb- 
ruary, and in the majority — in March. In young, estrus lasted from 
the second half of March to the first days of April. Parturition 
proceeded from the first days of April (old females) to 15-20 June. 
700 Duration of pregnancy ranged from 30-35 days. The number of 
embryos was from 2 to 6; average for adult females was 5.5 and 
for young — 3.6. Lactation period lasts 34-41 days. 

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 
litter 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 mor- 
tality (cause not established) was observed in 1941 (individual 
animals and litters) in Semipalatinsk district (Sludskii, 1963). 

Fluctuation in numbers of solongoi are well expressed. In 
periods of depression, their number decreases 2-4 times and after 
that, restored in the course of 2-3 years. From 1936-1949, num- 
bers of solongoi in Kazakhstan fell two-fold (Sludskii, 1953). In 
1939, the sharp decrease in its numbers coincided with a strong 
tularemia epizootic among rodents. Numbers of solongoi are also, 
without doubt, affected by the vast spring fires in the delta of the 
Hi river. In southeastern Trans-Baikaliya, the picture is different. 
Here, there were four increases in numbers of solongoi within a 9- 
year 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 without a black tip. It differs 
from the kolonok in its 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 
trapped. In the Balkhash region, the solongoi is harmful to com- 
mercial hunting, destroying muskrat and pheasant, and damaging 
the skins of muskrats in traps. Here, it may be considered a harm- 
ful predator. 

In agriculture and for ecosystem health, it may be considered 
a useful animal since 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 
annually (Stroganov, 1962) (P.Yu.). 

KOLONOK [SIBERIAN WEASEL] 

Mustela (Mustela) sibirica Pallas, 1773 

1773. Mustela sibirica. Pallas. Reise d. versch. Prov. d. Russ. 

Reiches, 2, p. 701. Tigeretsk Post near Ust-Kamenogorsk, 

Altai. 
1844. Mustela itatsi. Temminck. Fauna Japonica, Mamm., p. 34, t. 

VII, Fig. 2. Japan. 
1871. Putorius fontanieri. Milne-Edwards. Rech. Mamm., p. 205, 

pi. 61, Fig. 1. Pekin [Beijing], China. 
1904. Mustela sibirica miles. Barret-Hamilton. Ann. Mag. Nat. 

Hist. 13, p. 391. Dauriya, Trans-Baikaliya. 
\9\\. Mustela manchurica. Brass. Aus dem reiche der Pelze, p. 

490. Manchuria. 



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 con- 
striction is elongated, its lateral outlines almost parallel, no dis- 
tinct isthmus directly behind supraorbital processes. Dimensions 
relatively large — condylobasal length of male skull more than 55 
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 oc- 
cupy 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, one or several white small spots occur on 
throat. 
702 Tail usually brighter than back — bright-red above and below. 
It is covered by long, dense, loose-fitting hairs and is very fluffy, 
fluffier than all other species of genus, including polecats and 
minks. Tail thickest of all in middle part, gradually thinning 
towards its base and rapidly coming to naught at tip, which is 
tapered. Lower part of 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 fur, whole trunk covered by shorter, coarse, 
quite close-fitting hairs. On tail, hairs relatively short, close-fit- 
ting; therefore it 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 surround- 
ing 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 in color are absent. Young ani- 
mals are clothed in downy brown fur, 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 
(Zalesskii, 1970), white with light red guard hairs on back and tail, 
and several other mutational variants. "Giant kolonok" mentioned 
below have color which is not normal. 

The kolonok skull occupies, in several characters, an interme- 
diate position between the skulls of ermine and mink. It is larger 
and more elongated than ermine and is somewhat flattened com- 
pared 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 sig- 
nificantly less than distance from anterior edge of foramen 
magnum to anterior point of interpterygoid notch. Region of post- 
orbital constriction elongated, its isthmus almost unexpressed or 
weakly expressed, and outlines of lateral sides of this part of brain- 
case almost or more or less parallel. If something of an isthmus 
can be noted, then it is not situated close to supraorbital processes. 
Its 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 foramen less than alveolus of upper canine or equal 
to it. 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 between inner borders of anterior parts of bullae 
somewhat less than in posterior part. Distance between bullae in 
their anterior part less than width of hard palate at anterior edge 
of interpterygoidal notch. 

Zygomatic width approximately equal to skull width in mas- 
toid region or slightly more. Zygomatic arches elongated and weak. 
Protuberances, crests, etc. of skull relatively weakly developed, 
but generally stronger than in solongoi. Sagittal crest clearly 
noticeable in posterior part of cranium and in anterior, is weakly 
defined; occipital crest well developed. Upper profile of skull 
uneven; an elevation is noticeable on it in posterior part of 
cranium and prominence in frontal area (interorbital region) is ob- 
viously convex. From the frontal area, the line of the profile of the 
facial region 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 sex differences of the kolonok skull have not been 
especially studied. Generally, they correspond with those mentioned 
for other members of the genus Mustela (see description of er- 
mine). 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. Geo- 
graphic variation of the skull, within the boundaries of our coun- 
try, is not at all or is very weakly emphasized, and bears only a 
quantitative character (general measurements). 

The structure of the os penis is typical of the majority of 
species in the genus — the distal end is 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 
characteristic — more than in the species described above, com- 
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 
females, 49.8-62.7 mm; zygomatic width of male is 28.7-35.7 
mm, of females, 26.4-32.1 mm; interorbital width of males is 11.7- 
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 
(Stroganov, 1962), and also less — male (probably juvenile) is 395- 
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 
of the nose to the end of the tail with its terminal hairs reaches 
only 59-60 cm, then in giant animals, it ranges between 80 and 88 
cm. As well, color varies greatly — undercoat in "giant" kolonok is 
705 almost white (dirty yellowish-white or light-grayish), and paws and 
end of tail are dark-brown (Zalesskii, 1930; Zverev, 1931). There- 
fore, several characters of the steppe (white) ferret are well ex- 
pressed in them. Apparently, this is not an "exotypic" mutation of 
the kolonok, but a hybrid between kolonok and steppe ferret (M. 
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 illus- 
tration of heterosis, "Giant kolonok" are known only from skins 



^'From other data (Zverev, 1931; western Siberia), body length of males (55) is 
310^30 mm, often 350-370 mm, of females (60), 300-360 mm, often 260-310 mm; 
tail length of males (50) is 140-220 mm, often 160-210 mm, females (55), 130-210 
mm, often 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 (see section "Systematic Position" — 
in 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 between these 
species. 

The closeness of features of kolonok to mink are without doubt. 
In purely morphological features, the kolonok obviously differs 
from ferrets (subgenus Putorius); however, information concern- 
ing hybrids between these species occurring in nature (see above) 
point to a close connection between these seemingly separate spe- 
cies. This is, once more, confirmation of the homogenity of the 
genus Mustela, and at the same time an example of the compli- 
cated inter-relationships of its species. It is difficult to array them 
all 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 to the traditional one is adopted 
here as the most natural (V.H.). 

Geograpliic Distribution 

In Southeastern, Eastern and Central Asia, the southern and middle 
zone of Siberia and the Far East, and in the eastern European part 
of the USSR. 

Geographic Range in the Soviet 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 Komi district to the source 
of the tributary of lower Sukhona — the Luza (Oparino to the north- 
east, and Letka and Ob'yachevo to the north of Kirov), where it 
passes below 60°20' N. lat. (Ob'yachevo), and thence to the mid- 
dle Vetluga. From here, the border line apparently passes generally 
along the Vetluga and Volga to Kazan, and then is directed to the 
southeast, covering a great part of Tatariya, the northern- and 
northeasternmost outskirts of Kuibyshev district and Bashkiriya, 
and passes onto the southern Urals^°. 
707 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 kolonok range from бЗ** N. 
lat. in the Urals, crosses the Ob' river, passes to the upper Pur and 
then suddenly ascends to the north on the lower Taz, almost reach- 
ing the Arctic Circle here or, perhaps, even a little beyond it. To 
the east, the border still 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 then, returning to the east, goes 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 
middle course of the left tributary of the Vilyui — the Markha (64° 
N. lat.) and reaches the Lena to the north of the mouth of the 

'"According to V.A. Popov (Kazan), during the whole time of investigations in 
Tatariya, the kolonok was never found west of Chistopol'. This same author also 
considers that the animals which were caught in the Vetluga basin (Dryablovo, left 
tributary of the Vetluga — the Kaksha, 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. 




■^ >a 



u .^ 



••e «N 



1061 



Vilyui at 65° N. lat. Thence, it descends to the south along the 
right bank of the Lena valley, covering thereby the lower courses 
of its right tributaries and thus goes to the nnouth of the Aldan. 
Along the Aldan, also covering the lower courses of its right tribu- 
taries, the border goes to the Allakh-Yun' River and along it, ex- 
tends beyond the border of Yakutiya (D. Ivanov). From the 
Allakh-Yun', the boundary passes to the mouth of the Okhota at 
the Sea of Okhotsk (about 59°10' N. lat.; V.P. Sysoev). 

Individual cases of captured kolonok, apparently, dispersing 
individuals, were recorded in Yakutiya and somewhat northward 
of the above-mentioned line — along the Tyung in its upper course, 
along the Lena in the direction of Zhigansk and somewhat north- 
east of the mouth of the Aldan. 

South of the specified line, the kolonok is distributed through 
all 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 
Borzya, to Olovyannaya, and then to the left bank of the Onon, 
and thence southwest to the state frontier. In southern Altai, the 
kolonok extends south to the latitude of МагкакоГ Lake, but is 
absent in the Zaisan depression. 

To the west of the Altai, the kolonok is met with in the Kalbinsk 
Altai (left bank of the Irtysh), but farther south the border goes 
north again, generally along the Irtysh, although south of 
Semipalatinsk, the kolonok is 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 district. South of here, it penetrates to the area 
a little to the north (Alekseev forestry) and even a little 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 
Sakhalin, fossils of Siberian weasels were also absent, but in 1932, 
the Japanese weasel was introduced to this region, and from that 
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 
considerable changes and in a series of places these changes are 
continuing at present. In several places in central Siberia, Yakutiya 
and the Far East, the kolonok is extending its range, though not 
very intensively. Particularly notable is the kolonok' s colonization 
westward. Thus, even in the 70' s and 80' s of the last century it 
708 was absent in the Urals. The kolonok occupied the whole Euro- 
pean (Cis-Ural) part of its range, i.e. 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 it penetrated to the eastern parts of 
Gor'ki district (Vetluga basin); it was rarely encountered on the 
sources of the Luza — right tributary of the lower Sukhona, and by 
1949 it 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 
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 Perm district, 
lying within the border, it is 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] 
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 was found in all regions 
up to 1940, but in 1955, the southwestern 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 appears rarely, and transgressions do not occur every year, then 
becomes more common, inhabiting 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 is maximal, and includes 
also areas of transgression and greatest movements. 

Colonization was particularly intensive, first of all, in the Eu- 
ropean 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 de- 
creased. 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 
continuation of a process begun much earlier. This belief lacks 
supporting data, but it is probable that the kolonok colonized 
Siberia, or at least its more northern parts, in the past century. This 
is indirectly shown by the characteristics of its 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 no basis (Laptev, 1955)^^ for 
considering that the kolonok was generally absent in western 
Siberia in the 18th century and that it was only found east of the 
Yenisei (Bobrinskii, 1946). 

"This phenomenon is also well expressed in several other species, the southern 
distribution boundaries of which extend south to western Siberia and northern 
Kazakhstan. 

"The described range of the kolonok differs significantly from that which is 
usually described and illustrated (Bobrinskii, 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 
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 for the sake of acclimatization in 
1937 in the 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. 
The kolonok exists in the Mongolian Republic, where it occupies 




Fig. 253. Natural species range of kolonok, Mustela (Mustela) sibirica Pall. 

V.G. Heptner. 

*Contraction for urocliishche, meaning "small village" — Sci. Ed. 



1065 

northern montane region — Pri-Kosogol', Khangai and Kentei; on 
the Korean Peninsula and in China — northeastern (former Manchu- 
ria), 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**. 
710 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 con- 
sidered 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 de- 
mands further investigation, not only on the faunistics of Southeast 
Asia, but also the systematics of the kolonok (V.H.). 

Geographic Variation 

A series of forms have been described for kolonok, among them 
including 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 extend- 
ing from the Volga Basin to the Pacific Ocean, is very small — a 
small number of real forms and amplitude differences in characters 
is insignificant. This leads us to think that variation in the species 
is generally insignificant, and its presentation is exaggerated 
(Ellerman and Morrison-Scott, 1951). 

The scheme recently suggested for our territory (Stroganov, 
1962) is extremely fragmented and is mainly based in part on 
exceptionally subtle characters, the reality of which is highly doubt- 
ful. The recognition of the concept natio (for the form australis), 
which in essence is lacking in real content, does not make it more 
convincing. In exactly the same way, separation of the form itatsi 
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 in- 
sular forms, in particular itatsi are better distinguished* in natural 
conditions of occurrence outside our country. 

Within the borders of our country, two forms may be distin- 
guished. 

1. 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 
mm, 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, 
10.5-Ml 1.0-12.2 mm; mastoid width of males is 26.8-M27.5- 
28.7 mm, 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. 

Outside the USSR — found in the part of the Mongolian Re- 
public contiguous to its range and, possibly, in extreme western 
parts of northeastern 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 
(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, appar- 
ently, themselves represent a transition to the next form. All these 
deviations, even those in the Trans-Baikal population are, how- 
ever, weak and their territorial limits are not clear and do not offer 
a basis for separation of independent geographic forms. 

2. Far Eastern kolonok, M. (M.) s. manchurica Brass, 1911 (in 
our literature, the name coreana is accepted). 

Average measurements somewhat larger than in nominal form; 
color lighter, red. 
711 Condylobasal length of male skull (16) is 63. 8-M65. 4-67.3 

mm, 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; 
inter-orbital width of males is 12.3-M13.2-13.9 mm, of females, 

♦Contradictory? — Sci. Ed. 

"Measurement of this and the next forms from Stroganov (1962). Body measure- 
ments given above (p. 1057), from Zverev (1931), belong to this form. 



1067 

11.0-M12.7-13.0 mm; postorbital width of males is 13.2-M13.6- 
14.0 mm, of females, 12.0-M12. 7-13.3 mm; mastoidal width of 
males is 28.2-M29.5-31.0 mm; of females, 24.0-M25. 0-27.0 mm. 

This form is found in Priamur'e to the west to Zeya [river], 
Primor'e. 

Outside the USSR — in northeastern China (former Manchu- 
ria), precise limits unknown. 

A well defined form. Apparently, identical to the Manchurian 
[form] and must carry the name recognized here. Comparison of 
our Far East kolonok with Chinese and Korean, the names of which 
might be applied to ours (see synonymy) has not, however, been 
made and thus, the name manchurica, therefore, conditionally ap- 
plied. It is probable that the name fontanieri may be more proper. 

3. Japanese, or Sakhalin, kolonok^'*, M. (M.) s. itatsi Temminck, 
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 somewhat more 
bulging, inner border of auditory bullae somewhat separated 
posteriorly, and distance between auditory capsules in their poste- 
rior part is notably 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, 93- 
101 mm; length of hind foot of males is 57-65 mm, of females, 
39-46 mm. 

Condylobasal length of male skull is 57.2-61.9 mm, of fe- 
males, 47.6-57.9 mm; zygomatic width of males is 31.3-33.2 mm, 
of females, 28.9-32.0 mm; interorbital width of males is 12.0- 
12.8 mm, of females, 9.1-12.0 mm (after Stroganov, 1962; supple- 
mented by material of Z[oological] M[useum of] M[oscow] 
U[niversity]). 

Found in Sakhalin, only in south. Acclimatized there in 1932. 

Outside the USSR — in the Japanese Islands (Hokkaido, Honshu, 
Shikoku, Kyushu, Iki). 

'^It was suggested (Pereleshin, 1957; Stroganov, 1959, 1962) that this form be 
given the 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, 
especially a non-declinabie one. 



1068 

A well-distinguished form, differing more strongly from named 
forms than the latter do from each other. 

Note. The relatively sharp differences between Japanese kolonok 
and Siberia turned attention to it when first recorded within the 
borders of the USSR (Pereleshin, 1957). 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 this author and the majority of characters noted by him 
(12 in all), in particular, 5 out of 9 of which are craniological, a 
"sharp hiatus" exists between the form itatsi and those of the 
mainland. 

Materials in the Z[oological] M[useum of] M[oscow] 
U[niversity] from Sakhalin (collected by A.I. Gizenko), however, 
show that the features distinguishing the form itatsi from Siberian 
are far from such as those described and illustrated by Stroganov 
(1960, 1962). A clear hiatus between is 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 audi- 
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 its length in sibirica) 
as well as dimensions and form of the bulla — are not demonstrated 
712 at all in material of ZMMU. The same applies to other [charac- 
ters]. Some bulging of the frontal area is noticeable (weak) and a 
well expressed difference in the position of the inner outline of the 
boney auditory bulla, as mentioned in the diagnosis above. 
Therefore, to believe that the form itatsi inhabiting Sakhalin is an 
independent species is without foundation. It is a true kolonok and 
its somewhat greater differences are, apparently, explained by its 
insular origin. 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.) s. subhemachalana Hodgson, 1837 — Himalayas from 
Nepal to Bhutan; 2) M. (M.) s. canigula Hodgson, 1942 — Tibet; 3) 
M. (M.) s. hodgsoni Gray, 1843 — Kashmir and western Himalayas 
from Kam to Garwal (Garhwal); 4) M. (M.) s. davidiana Milne- 
Edwards, 1871 — Southeast China north to Hubei, Taiwan; 
5) M. (M.) s. fontanieri Milne-Edwards, 1871 — Northern China, 
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. 
(M.) s. coreana Domaniewski, 1926 — Korean Peninsula; 9) M. (M.) 
s. charbinensis Loukashkin, 1934 — Manchuria; 10) M. (M.) s. 
quelpartis Thomas, 1908 — Quelpart Island in East China Sea; 11) 
M. (M.) s. sho Kuroda, 1924 — Tanageshima and Yakushima is- 
lands (southern Japanese archipelago) [= Ryukyu]; 12) M. (M.) s. 
asaii Kuroda, 1943 — Oshima Island, Japan. 

The list given is obviously inflated, probably approximately 
twice. For some forms, deviations in characters of the facial 
pattern, the majority of which are 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 diver- 
sity 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 latter produces more than 1/3 of the skins prepared in 
the USSR. In the Gorno-Altai Autonomous District, the highest 
frequency of tracks found (in 10 km) in a series of years ranged 
from 2.85 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 
a 10-km route ranged from 24.1 to 38.8 in half the cases, but 
sometimes it decreased to 2.1 (Yu.A. Salmin and V.D. Shamykin). 
Population and distribution of kolonok are determined by food 
abundance (mouse-like rodents and water voles), the stability of 
stocks, interrelationships with competitors (in dark coniferous 
taiga — with sable; in taiga river valleys — with ermine and also 



1070 

American mink; at the upper edges of forests — solongoi and 
ermine; at the steppe border — with steppe polecats) and also the 
characteristics of snow cover. 

Habitat. Kolonok habitats are varied. In the taiga, it willingly 
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, after old burns, dense growth of saplings. 

In the Altai mountains, the kolonok prefers dark coniferous 
taiga with larch stands. However, these are secondary habitats 
whence it penetrates after the extermination of sable, and from 
which after a time sable again 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 



Fig. 254. Mountain creek — typical habitat of kolonok. "Kedrovaya Pad' " preserve, 
southern Primor'e. October 1963. Photograph A.G. Pankrat'ev. 



1071 

713 type and in coastal oak forests. It is met with from the sea coast 
to the 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" 
along the banks of steppe lakes and rivers, and meadow steppes 
with shrubby growth, and rarely — pine forest islands. It 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 
the other causes damage to domestic fowl. It 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 (% of occurrence) 











л о 


a 


> 












JS. i! 




о 


я 




> 
о ^ 


00 

<u . 


i» en 


5S ^ 
2 § 


ее 
N 


stem 
(Fetis 
2) 


N^^ TO 


Type of food 


55 > 


ppe, w 
Zverev 
931) 


« 2 
.2 '^ 


ern Al 
, 1956 
62) 


^ 

a^ 

S 2 


ppe,we; 
kaliya 
7, 194 






C/3 о 

< (U 


1) ■-^ -< 
? =« > 




1 °- 


^ 2 


a -5 g 

1Л и _4 






- H 


tS •= 2 


я 'С 


•Й Й Й 


^^ ^ v^" 


M -s 




CQ 


(U u E 


ш u 


t; ■= о 


XI > 


У '=' iS 


.2? ^ 




■^ -o 


I- X) •- 


•^ .Q 


о '" fl 


r^ ce 


u. (Я ro 






ra a 


о .- — ; 


Я •- 


►2 -J 


о ,t- 0\ 


' я "ее 




H rt 


[1, CO t^ 


H 00 


Z CL OX) 


z. d- 


11- H - 


Н с/5 


Water vole [Arvicola 


36.6 


56.1 


46.0 


1.8 


7.0 


— 


— 


terrestris] 
















Small voles and mice 


15.0 


15.6 


15.0 


50.1 


20.0 


24.6 


68.7 


Altai zokor 


— 


— 


— 


— 


21.0 


— 


— 


[M. myospalax] 
















Great jerboa 


— 


— 


— 


— 


14.0 


— 


— 


[Allactaga major] 
















Siberian chipmunk 


— 


— 


— 


0.9 


— 


9.6 


— 


[Tamias sibiricus] 
















Pikas 


— 


— 


— 


2.7 


— 


38.2 


— 


Insectivores 


6.6 


0.8 


— 


33.8 


2.0 


3.5 


— 


Birds 


3.3 


4.6 


15.0 


6.4 


29.0 


13.6 


9.4 


Fish 


13.3 


— 


15.0 


— 


— 


0.5 


4.4 


Carrion 


— 


1.7 


— 


— 


— 


— 


10.1 


Amount of data 


60 


130 


43 


108 


52 


406 


684 


examined 

















1072 



kolonok food everywhere. Species composition and ratio of each 
species depend upon the local conditions and competitive relation- 
ships with other carnivores. Geographic variation in primary 





714 Fig. 255. Bank of a montane creek — characteristic winter habitat of kolonok. 
"Kedrovaya Pad'" preserve, southern Primor'e. March 1964. Photograph 

A.G. Pankraev. 



(occurrence from 15% of cases) and secondary (occurrence from 
5%) of foods is shown in Table 67 for seven different regions in the 
range. In it are placed only those foods and groups were given which 
were found, at least in 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 number of main foods include ro- 
dents of moderate size — Daurian and 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 some places. In Ussuri 

*In the Russian original, this word is spelled "Alaiskaya", but this region is west 
of the one being discussed, and "Altaiskaya" was probably meant — Sci. Ed. 



1073 

territory, carrion — food remains from wolf and yellow-throated 
marten — are very important in winter. Everywhere, birds play an 
essential role in food of kolonok, chiefly small ones — to a lesser 
extent, poultry. Attacks on domestic birds are a widespread oc- 
currence, as is damage to trapped hares, squirrels, kolonok and 
other animals. Feeding on reptiles and amphibians is observed at 
the periphery of the range — in Tatariya and especially in Primor'e, 
Among a number of plant foods are found nut pine, "nutlets", 
fruits of actinidia vines and others. However, in the montane taiga 

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 
rodents. Carrion is eaten from November to March; birds — in June- 
August, and fish — in September-October (migration of salmonids). 

715 The role of plant food (not above 4.9%) increases in August — 
February. Amphibians, reptiles and invertebrates are foods of the 
warm period of the year. The frequent occurrence of pikas (25%) 
is 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 to 12.4%, plant foods— from 1.0 to 
8.0%, etc. (food was studied in 12 habitats; Yu.A. Salmin and 
V.D. Shamykin). 

The proportions of other foods change from year to year. In 
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 8-10 km. In the northeastern Altai, the area of 
daily activity constitutes 1.5-2.0 km^ while the home range is equal 
to 4-5 km^. 



1074 

Burrows and shelters. Shelter chosen by the kolonok is 
varied. It may be a 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 deepen- 
ing them. In western Trans-Baikaliya, the length of burrow pas- 
sages of kolonok appear to be from 0.6 to 4.2 m, and depth — from 
0.2 to 1.3 m. The nesting chamber is located in the middle or at 
the end of the passage, and is 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), 

Daily activity and behavior. Crepuscular and nocturnal activ- 
ity is typical of kolonok, but its rhythm is not clearly expressed — 
diurnal activity of kolonok is a quite common phenomenon, 
especially in summer and autumn. It hunts in the morning and 
during the day, as well as in winter when severe frost occurs at 
night and during the day warms up (Shaposhnikov, 1956). During 
times of severe frost and snow-storms, the kolonok does not ven- 
ture 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) — to 10 km and more, and 
in the northeastern Altai — from 1.5-2.0 to 6-7 km. Its length de- 
pends on the quantity of prey and condition of snow cover 
(Shaposhnikov, 1956). In contrast to sable, in winter the kolonok 
hunts food more actively, mainly under the snow — under logs, in 
windfalls and beneath the arches of snow-laden bushes while sable 
more often lies in wait like a cat. Compacted snow and frozen 
crust makes hunting under the snow almost impossible for the 
kolonok, and therefore, it confines itself to the northeastern slopes 
of montane taiga in the Altai, and terraces of the northern quad- 
rant, where the snow is always loose and where frozen crust does 
not form (P.B. Yurgenson). During one night, the kolonok enters 
into the snow more than 30 times during a distance of 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 



716 



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 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. Tracks of a young male kolonok on soft first snow on the ice of a mountain 
creek. "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 in 1928 are more 
reliable. The massive transfers were observed in a northern direc- 
tion, 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. 
Seasonal migrations occur in connection with changes in condi- 
tions of food capture and their dispersion among habitats. 

Reproduction. In the kolonok, the period of rut is quite ex- 
tended. 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 
continues to the end of April. In the Pushkin fur sovkhoz near 
Moscow, rut in six pairs of kolonok took place from 25 April to 
15 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 accord- 
ing to other data — within limits of 35-42 days (Kler, 1941). Par- 
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 al- 
most 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. 

Molt occurs twice a year — in spring and autumn. 

718 Enemies, diseases, parasites, mortality, competitors, and popu- 
lation dynamics. Sable, both species of ferrets, red fox and eagle 
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 unknown reasons are known 
(Sludskii, 1963). 

Parasitic infection of kolonok, according to the data of Sikhote- 
Alin' preserve (Yu.A. Salmin and V.D. Shamykin), is very 
considerable. In 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 10- 
15 to 30 individuals). Infection with other endoparasites is low. 



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 
increase in numbers of kolonok follow years of massive rodent 
reproduction. For some regions of western Siberia, increases in 
numbers of kolonok were recorded in 1910, 1916-1917, 1922- 
1925 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 
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 ani- 
mal is 35 cm; a frightened kolonok makes leaps of up to 1 m. In 
contrast to ferrets, running of the kolonok has a somewhat con- 
fused character, is twisted, and more closely approaches that of 
mink, ermine and weasel (P.Yu.). 

Practical Significance 

The kolonok is a valuable fur animal, having a significant share of 
fur preparation of Siberia and the Far East. The fur of the kolonok 
is used as both natural and for imitating more valuable species. 
The guard hairs of the tail are highly valued: used in the prepara- 
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, p. 5. Finland. 
1777. Lutra minor. Erxleben. Syst. Regni Anim., 1, p. 451. Re- 
naming of lutreola Linnaeus. 



1079 

1879. Lutreola europaea Homeyer. Zoolog. Garten, 20, p. 184. 

Renaming of lutreola Linnaeus. 
1912. Mustela {Lutreola) lutreola wyborgensis Matschie. 

Sitzungsber. Ges. naturforsch. Freunde Berlin. 1912, p. 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 district; V.H.). 

719 1912. Mustela (Lutreola) lutreola budina. Matschie. Ibidem, p. 

349. "Ortelsburg, East Prussia" (Shchitno southeast of 
Ol'shtyn [Olsztyn], northern Poland; V.H.). 

1912. Mustela (Lutreola) lutreola varina. Matschie. Ibidem, p. 351. 
Vic. of Shverin, Mecklenburg. 

1912. Mustela (Lutreola) lutreola albica. Matschie. Ibidem, p. 351. 
Levitts [Lewitz] river — tributary of the Elbe. Mecklenburg. 

1912. Mustela (Mustela) lutreola glogeri. Matschie. Ibidem, p. 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 — about 150 km southwest of 
Krakow; V.H.). 

1932. Mustela lutreola hungarica, Ehik. Ibidem, p. 142. Kovachna; 
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 — Mustela flavigula var. borealis Radde, 
1862. 

1939. Lutreola lutreola caucasica. Novikov. Ibidem, p. 63. 
Prokhladnaya Station, northern Caucasus. Nomen praeoc- 
cupatum — 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 
neck. Postorbital constriction of skull weak — its width equal to or 
more than width of interorbital distance. 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.). 

Description 

Concerning general appearance, the mink is a typical representa- 
tive 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 to 
ferrets, greatly reminiscent of the black (M. putorius). 

Head in mink relatively quite large, with broad, but not tall, 
ears (in winter, they slightly protrude from fur). Limbs short, 
relatively well developed membranes between digits, particularly 
on hind paws. These are larger than in ferrets and only terminal 
720 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 less similar, close fitting hairs similar in length through- 
out, 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 
quite loosely fitting. Compared to terrestrial species of the genus, 
underfur particularly dense. Guard hairs quite coarse, lustrous, and 
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 life. 

Summer fur somewhat shorter, coarser and less dense than 
winter one; however, this difference is 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 
cases, ventral surface is 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 
some even black, broad dorsal belt is noticed. Limbs are slightly 
darker than trunk, as is tail, the end of which may be almost black. 
Underfur is bluish-gray. Mink fur, particularly dark individuals, is 
very beautiful and has a dense silky luster due to dark guard hairs. 

Mink has no color pattern at all on the head, and it has a color 
in common with the whole trunk, but upper and lower lips and 
chin are pure white. Pattern of upper lip is quite stable, but on 
chin, varies in size and form. It 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 
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 
721 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 
occupies the throat region; with this there may be a spot or spots 
on the chest. Finally, and not too rarely, in some places the whole 
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 
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 some quantity of them spreading 
throughout the pelage are completely white. Extreme exotypic 
deviations in the form of albino mink, or pelage covered with 
separate white spots, 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 — without dark silky 
or silvery luster. 

On the whole, individual variation in color, if white markings 
on chest and throat are not considered, is not great. Sexual differ- 
ences 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 is relatively small (see below) and is 
manifested in lighter and more reddish color of fur from north to 
south. White markings on throat and chest are quite rare in the 
north, and are encountered more frequent in a southerly direction; 
their general area increases and they attain their greatest develop- 
ment in the extreme south of the range (Caucasus). Just here — and 
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 sev- 
eral 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 rest, the mink skull is characterized by the following 
characters. It is 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 weak — its width at the 
narrowest point is approximately equal to or greater than the inter- 
orbital width. The zygomatic arches are relatively weak and are 
moderately widely separated, the mastoid processes are relatively 
weakly developed. The auditory bullae are small, swollen, rela- 
tively short and narrow, and widely separated; in the posterior 
part, they separated from each other significantly farther than in 
the anterior part. The ratio of their length to their width is approxi- 
mately 1 : 1.7. Sculpturing of the skull is not sharp, crests and 
prominences are moderately developed. The occipital crest is well 
defined, the sagittal one is 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 is not greater or only slightly greater 
than the longitudinal diameter of the outer. The posterior end of 
the second upper premolar is in contact with the anterior outer 



1084 

edge of the carnassial tooth, but it does not lie in the groove on 
723 its anterior edge. The inner surface of the 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 less 
than in the American mink. 

Except for dimensions (see below) the female skull is differ- 
entiated by some less developed protuberances, crests, etc. and 
lower weight, as in all other members of the genus. The juvenile 
skull is characterized by weaker development of the facial part, its 
shortness and a more swollen braincase. 

Geographic variation of the skull, besides general dimensions, 
several of which exhibit growth from north to south, is manifested 
in the fact that in the extreme south (Caucasus) the elevation of the 
skull in the interorbital region increases (strengthening its similar- 
ity to the kolonok) and several features appear (greater constric- 
tion of the postorbital region), 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 major- 
ity 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 
the lower side of the bone 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 
somewhat to the right. 

According to body and skull measurements, the mink belongs 
to the group of large species of the genus — the ferrets and kolonok. 
Dimensions of the species as a whole: body length of males is 
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, 50- 
M52-53 mm; height of ear in males is 20-M23-25 mm. Male tail 
constitutes 35-M41-52% of body length, of females, 37-M40- 
45%. 

Condylobasal length of male skull is 56.4-M62.7-68 mm, of 
females, 52.8-M61.4-65.7 mm; interorbital width of males is 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-M32.3- 
37.0 mm; mastoid width of males is 27.0-M31.7-34.9 mm, of 
females, 26.5-M29.4-31.7 mm (Novikov, 1939). 



1085 






;У>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-M327- 
360 mm; tail length of males is Ml 45 mm,* of females, 120- 
M128-142 mm; length of hind foot of males is 52-M56.2-61 mm, 
of females, 45-M50.0-58 mm. 

Condylobasal length of male skull is M67.5 ± 0.40 mm, of 
females, M63.7 ± 0.52 mm; zygomatic width of males is M35.2 ± 
0.52 mm, of females, M30.5 ± 0.65 mm; interorbital width of 
males is 12.9 ± 0.30, 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, 
of young, (10) 32.6, 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. 
Popov, 1949). 

The total weight (from data for Middle Europe) is 550 to 800 
gm (V.H.). 

Systematic Position^^ 

The systematic position of mink among those species which are 
here united in the genus Mustela, has been evaluated entirely dif- 
ferently by different authors. An extreme point of view holds that 
the 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 
over-estimation, of a known hypothesis of their unique way of life. 
724 Right now, this point of view has almost no supporters. More 
often, both species were included, and are included, in the genus 
Mustela in the category of a separate subgenus, equivalent to fer- 
rets. Isolated opinion holds that minks represent a special subgenus 
in the 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 rep- 
resents a development and combination of the same set of features 
which characterize the previously described species of the genus. 

*No range given — Sci. Ed. 

"For section on American mink, see end of book in "Supplement to the order of 
carnivores". 



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" position of mink is emphasized by the fact 
that in nature hybrids between mink and black ferret are encoun- 
tered, and these do not represent exceptional rarities. There are 
such specimens in museums (Z[oological] M[useum of] 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 
skins of hybrids, characteristics of one or the other parental spe- 
cies predominate (Kuznetsov, 1952). Remarks stating that "data on 
crosses between mink and ferret are completely absent (Novikov, 
1939) is based on misunderstanding. Nothing is known concerning 
fertility of "khor'-tumak". 

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 morphologi- 
cal characteristics which divide them are substantial. Hybridiza- 
tion of both species in nature, which is sometimes assumed for 
Central Europe (Gaffrey, 1961), is not confirmed. On the contrary, 
in 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 to cross both species in captivity were not successful. 

In the sense of the structure and phylogenesis of the genus, it 
is 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 — at the present 
time, nearly the whole. It occupies the European part of the Union, 
Caucasus and the western part of western Siberia. 

The northern border of the range in the extreme northwest — 
in Karelia, passes a little north of the Kem' river — at the southern 
extremity of Topozera [lake] and Kuzema [city] (Marvin, 1950, 
725 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 line, passing south around the Kanin 
tundra, adjacent to Cheshsk inlet and Malozemel'skaya tundra, 
passes around Sula (left tributary of lower Pechora) from the north, 
and along the Pechora, extends as a projection, to the delta, i.e. to 
68° N. lat. (Novikov, 1939; V.Ya. Parovshchikov). This is the 
northernmost point of the range. 

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, therefore, to the southern part of 
the forest-tundra or along its 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 somewhere around the Arctic Circle or 
probably south of it^'. Farther, the border goes directly south, in- 
cluding the Lyapin river (noted mainly at Saranpaul') and the 
upper course of Severnaya Sos'va. Thence, in an as yet undeter- 
mined path, directs itself, apparently, again southwards and then 
turns eastwards reaching the Ob' somewhere a little 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 tribu- 
tary of the Ob' (I. Laptev; perhaps transients). Thence, the border 
line goes directly south, covering the basin of the Dem'yanka (mink 
is known from several places in its middle course) and reaches 
Тага on the Irtysh. Beyond the Irtysh, the range does not extend 
above its course (Yanushevich and Arkhangel'skii, 1952; I. Laptev, 
1958). 

The southeastern and southern borders of the mink range in 
western Siberia is poorly known. On the whole, from Тага, it arches 
to the Ural ranges, somewhere approximately at the latitude of 
Chelyabinsk. South-southwest of Тага the border, apparently, passes 
a little north of Petropavlovsk (transient mink are known at Lake 
Chagly north of Kokchetav; Sludskii, 1953) and at Zverinoglovskaya 
on the Tobol south of Kurgan (I. Laptev, 1958). 

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 
reaches the Ural at Ural'sk or slightly to its south. Mink do not 
occur lower along the Ural [river]. The suggestion (Novikov, 1939) 
that mink is absent in the southern Urals and inhabits only the 
western slope in this region, is incorrect (Shvarts, Pavlinin and 
Danilov, 1951; Kirikov, 1952). Probably transient mink are known 
at Ilek, on the Irgiz river (Кора settlements; Sludskii, 1953). Infor- 
mation concerning the occurrence of mink in Alakor(Semirech'e; 
Shnitnikov, 1936) are lacking any basis. 

From Ural'sk the border, going west, extends over the basin of 
the Volga tributaries — Bol'shoi Irgiz (Elpat'evskii, Larina and 
Golikova, 1950; mink are absent on the Uzen') and descending 

"The reference that the mink is distributed in the Urals nearly to 70° N. lat. 
(Shvarts, Pavlinin and Danilov, 1951) was based on some sort of misunderstanding, 
or it is a misprint; 70° N is 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- 
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 
727 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 to the Sal, reaching its lower course. 
Thence, it 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 Kuma basin, also including its northernmost tribu- 
taries. Along the Kuma, the range border runs to the sea, or some- 
what short of it. 

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 at the source of the Kuban), it goes 
in an irregular line through the upper reaches and sources of the 
Terek, Kuma and Kuban river systems — approximately to the 




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 
Black and Azov seas (present in Taman'). In the east, between 
Deshlagar and the mouth of the Terek, the range is 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 Cau- 
casus, 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 who is known for his ruinous errors. Generally 
speaking, little is known on the course of colonization in the mink 
(for details, see Novikov, 1939; I. Laptev, 1958). That it is con- 
tinuing to settle new places at the present is not excluded. This is 
728 shown by the above-mentioned transgressions of mink to Kokchetav 
and the Irgiz. Beyond the Irtysh (upper Тага), however, the mink 
was still absent in 1950 and 1960 (for details, see Novikov, 1939). 

'«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 this, there was a suggestion recently, not lacking 
in serious foundation, speculating that mink in western Siberia 
occurred very long ago — even before the academic expedition in 
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 mink in western Siberia, there is a situation analo- 
gous to that previously mentioned for the pine marten (see p. 844)*. 



? 



0] 




. *<:ь 



е-гГ 



У 



zr 





Fig. 261. Species range of the Russian mink, Mustela (Mustela) lutreola L. In Europe, 

beyond the borders of the USSR, the range is 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 is absent concerning colonization of mink in other 
directions. Some changes in range are to be expected in connec- 
tion with the introduction of American mink in many parts of our 
country. 

Geographic Range outside the Soviet Union 

This (as established in the 19th and 20th centuries) includes Fin- 
land northwards to 66° N. lat. or a little farther, Poland, Hungary, 
729 Czechoslovakia, Romania, [eastern]* Austria, Bulgaria north of 
the Danube and Dobruja**, northern and western Yugoslavia — 
Istria, Croatia, Bosnia and Herzegovina — i.e. in the west of those 
countries south to approximately 43° N. lat. and in the east, to 
Belgrade, the territory of the German Democratic Republic, a con- 
siderable 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 beyond the bound- 
ary of our country and its range is shrinking. It has already disap- 
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 in western Europe apparently 
lies chiefly in its destruction by the American [mink] (V.H.). 

I Geographic Variation 

A quite large number of races have been described from the 
comparatively small species range of the mink, even if we do not 

♦Russian original ambiguous — Sci. Ed. 

♦*Not clear; Bulgaria lies entirely south of the Danube, and Dobruja is 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, 
now 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 Yugo- 
slavia were given by B.M. Petrov. References encountered in the literature concerning 
the previous presence of mink in the Scandinavian Peninsula and its disappearance 
not too long ago and its replacement by the American mink (crowding it out or its 
absorption by hybridization), are incorrect. It was never present there (see Ekman, 
1922). 



1095 

take into consideration the notorious synonyms given by Matschie 
(1912). In actuality, geographic variation in the species, first of all 
in the 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 
with water is 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 comes from misunderstanding (Ellerman and Morrison- 
Scott, 1951, of the Caucasian 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 increase, fur lightens, 
coarsens and becomes less dense. On the whole, both these char- 
acteristics, 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] M[useum of] M[oscow] U[niversity]). 

1. 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, bluish- 
gray. Pelage long, compact and silky. 

Body length of adult males is usually 340-M365-380 mm; tail 
length is 124-M 137-1 50 mm; length of hind foot is 54-M56-58 
mm; tail length is 36-42*, averaging 38% of body length. Weight 
of 100 skins is 6.5 kg. 

Condylobasal length of male skull is 56.4-M61. 5-64.3 mm, of 
females, 52. 8-M58. 2-64.3 mm; interorbital width of males is 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-M32.1- 
37.0 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'kii- 
Kazan'-Sverdlovsk; possibly in Bashkiriya. 

Outside the USSR — in Finland. 
730 2. Middle Russian mink, M. (M.) /. novikovi Ellermann et 
Morrison-Scott, 1951 (syn. borealis). 

Body and skull dimensions moderate — somewhat larger than 
preceding form. 

Color somewhat lighter than in 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 11.9- 
M13. 8-14.7 mm, of females, 11.3-M12. 3-13.9 mm; zygomatic 
width of males is 30.8-M36.3-38.6 mm, of females, 31.2-M33.0- 
36.7 mm; mastoid width of males is 28.4-M3 1.2-33.2 mm, of 
females, 28.0-M29.2-30.3 mm. 

Found in middle zone of European part of Union, south of 
distribution area of previous form in west, including Estonia, east- 
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 in Bashkiriya is not clear. 

Outside the USSR, probably not encountered. 

3. Caucasian mink, M. (M.) I. turovi Kusnetzov [in Novikov]*, 
1939 (syn. caucasica, binominata) . 

Body and skull dimensions large — the largest of the races liv- 
ing within the boundary of the USSR. Skull relatively massive 
with well developed protuberances, crests, etc. and strong zygo- 
matic 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 (red- 
dish) highlights. Underfur light, bluish-gray. White markings on 
chest frequent, and much more often than in other forms; in many 

*See note below — 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 
skins about 9 kg. 

Condylobasal length of male skull is 60.8*-M58.2-68.0 mm, 
of females, 54. 0-M56. 3-59.4 mm; interorbital width of males is 
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-M33.3- 
35.8 mm; mastoid width of males is 29.4-M3 1.8-34. 9 mm, of 
females, 30.5-M30.8-31.2 mm. 

In 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 
as its synonym. Later, N.A. Bobrinskii (1944) applied the name 
"turovi Kuznetzov et Novikov, 1939" to the Caucasian mink. In 
731 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 oc- 
currence (see above, synonymy of species); the Caucasian form of 
the mink must be named as given above, but not attributed to G.A. 
Novikov or to both authors. 

Of the three described forms, the most distinguishable is the 
northern, M. (M.) I. lutreola and the Caucasian, M. (M.) /. turovi. 
They are well delineated forms differentiated both by their body 
and skull dimensions, and also fur color and characteristics, and by 



*Sic\ judging from values of mean and maximum, probably should be 50.8 mm — 
Sci. Ed. 



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, 
M. (M.) I. turovi. Color quite dark, apparently, corresponding to 
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 13.3- 
M14.2-15.2 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; mas- 
toid 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: 
Poland. 

A very little known, essentially doubtful form, the character- 
istics and distribution of which are poorly studied in our country 
and which cannot be considered established; the same, it can be 
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 only possible now on the basis of material from our 

■""The name hungarica is considered to be a synonym of the form transsylvanica. 
As shown above (see synonymy), it is described from the immediate vicinity of the 
type locality of the form gloyeri, i.e. cylipena. 



1099 

territory. It is not excluded that the Middle European form is very 
close, 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 dark- 
tawny. 

Found in Moldavia (?) and Carpathia.'*' 

Outside the USSR — in Rumania, possibly Hungary, Bulgaria 
and Yugoslavia. 

To the highest degree, a poorly known and entirely doubtful form. 
732 Of the described forms having a relationship to our fauna, 
only two or three (nominal, Caucasian and Middle Russian) may 
actually be considered real; the remainder are very doubtful and 
basic study 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 USSR, still another form is usually 
recognized, M. (M.) /. biedermanni Matschie, 1912 — France (V.H.). 

Biology 

Population. The distribution of the mink within its range has the 
character of a fine lacey network, since its innate area is restricted 
to a 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. On the latter is depend- 
ent the density of 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 — Sci. ed. 

■"According to some data (Konyukhovich, 1953), the mink, which is quite 
widely distributed in Carpathia (Zakarpatsk district), is relatively large and dark 
and differs from the Middle Russian form. It is now impossible to decide from 
these data whether it is the given form or the Middle European cylipena. 



1100 

country, 24% of mink were captured, in Arkhangel'sk and Vologod 
districts and in Komi ASSR — 25%, in the Volga area, Bashkiriya 
and Urals — 23%, and in Kareliya and Leningrad district — 15%. 

If, 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, it became two — three times less. The causes 
of this decrease are several: a general decrease in intensity of the 
trapping industry, its crowding out from a series of areas by the 
American mink, local reduction in 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. 

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 woody banks as regards food abundance and the pro- 
tective conditions. Nevertheless, mink are met with on such rivers 
as the Volga, Kama, and Vyatka (V. Popov). Mink has not mas- 
tered the water surface — it is an inhabitant of the riparian zone. 

The dimensions and depth of the water in the 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 sec- 
tions — polynyas, broken ice, springs — are very important places. 
No less important is freezing of the water at a high level, with 
formation then, when it recedes, of a considerable space below the 
ice. Total freezing of the water is very unfavorable, but when 
winter food is abundant or when accumulations of hibernating frogs 
are available, winter survival is not inhibited. 

Water bodies where "naled" are formed [water on top of ice 
surface] are unsuitable for mink. With incomplete freezing, mink 
feel themself in excellent conditions in winter in very small bodies 
733 of water — 1.0-1.5 m 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 



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 are extremely variable as re- 
gards feed conditions, denning and protection. All of the variation 
of water bodies as denning areas, 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 all 
elements of the aquatic and riparian fauna. Its food varies depend- 
ing on composition of this fauna, time of the year, crops of differ- 
ent 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 in the European parts of the Union. 

The daily food requirement of the mink food consists of 140- 
180 gm. When food is abundant, mink makes stores. Usually, they 
are not great and contain, for example, up to 20 loaches 
[Miagurnus], 3 large ide and pike or several water voles. Sometimes, 
the store consists of frogs alone. When there are outbreaks of fish 
kills, 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 
water voles was twice that of the preceding year (14.2 and 33.3%). 
On 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 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 
plentiful and this food is more calorific than fish and frogs. 

Home range. The range inhabited by individual animals is not 
large, possibly thanks to the generally high food resources on the 
banks of small water bodies. The dimensions of individual home 
ranges fluctuate depending on feed resources of a given region. In 
regions of water meadows with little food, the home range attains 
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 fluc- 
tuates 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, in- 
dividual 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 
734 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. The burrow is used 
throughout the year, with the exception of the period of the spring 
freshets and summer floods. Usually, the burrow is located not 
more than 6-10 m from the water's edge. It is of simple construc- 
tion: 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- 
times, a lining does not occur at all. 

Often, the burrow opens to the outside under water, as in the 
river beaver, otter and desman. Such underwater outlets are con- 
structed when the bank of the water body, though not high, is 
abrupt and steep. In other cases, the burrow is made under the 



1103 



roots of trees, while in marshy places, the burrow is made in high 
hummocks — "kobla" of European alder or spruce. Often, mink set- 
tle in the hollows of riparian trees 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 
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 availabil- 
ity 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 some- 
times, during the transition from completely frozen bodies of water 
to those conditions more favorable, and in spring during the breed- 
ing season. 

The greater part of the time, mink go about on the shores of 
water bodies. Except for hours of rest, it finds itself in continuous 
movement in search of food. All its movements are quick and 
bustling. In case of danger, it tries to hide itself in water, in ex- 
treme cases, it conceals itself under roots or fallen trees. It 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 2-3 seconds and dives again. The mink climbs 
poorly, but in pursuit, it can climb a tree up to a height of four m. 
It hides well and observations of it rarely present themselves. 

The mink is an animal in which sedentariness is well devel- 
oped. Due to this, within separate bodies of water, it is very easily 
killed out. It 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, lim- 
ited 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 in search of 
females. At that time, paths of minks are observed along the banks 
of rivers of Moscow district (P.B. Yurgenson). 
735 Reproduction. In Moscow Zoo, estrus was observed on 22-26 

ApriH'. 

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 Euro- 
pean 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 20- 
25 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 popu- 
lation 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 number of otters increases the 

*H.e., approximately one month later than in the American mink. It is supposed 
(Rubetskaya et al., 1933) that this is 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 it to its burrow (Formozov, 1923). 
736 Diseases of minks in nature have not been studied. Invasive- 
ness with helminth worms was determined as 56% (M.P. Lyubimov). 
Among helminths, 27 species were recorded in minks: trematodes — 
14, cestodes — 2, and nematodes — 11 (Petrov, 1941). Particularly 
often, pulmonary filariasis and krenzomatiasis are encountered, as 
well as skryabingulosis infecting the frontal sinuses. They some- 
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. Neya river, Ponazyrevsk 
region, Kostromsk district. 22 October 1939. Sketch by A.N. Formozov, about 2/3 

natural size. 



1106 



Mink populations independently of the influence of those 
harvested do not remain at one level. However, a marked sequence 
in its variation has not been established. Usually, populations grow 
for 3-5 years. Such widespread death of mink is scarcely con- 
nected 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 mouse- 
like 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 restless- 
ness 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 in that period when the rising water fills all the empty 
places under the ice and drives the mink to dry land. Spring hunt- 
ing is 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. 

n 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. Mamm. Brit. Mus. Sub- 
stitute for putorius Linnaeus. 

1851. Putorius verus. Brandt. In: Semashko. Russian Fauna, 2, p. 
357. Substitute for Mustela putorius Linnaeus. 

1926. Putorius putorius stantschinskii. Melander. Nauch. izvest. 
Smolensk, gos. univers., p. 137. Roslavl'sk со., Smolensk 
gov. Described as a color "variety". 
737 1929. Putorius putorius orientalis. Brauner. Ukrainsk. mislivets 
ta. Ribalka, No. 2-3, p. 9, Ukraina. Exact type locality not 
indicated, but apparently, northern part of former Khersonsk 
gov. Nom. praeocc. — 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 
part of USSR. Nomen praeoccupatum — Mustela erminea 
ognevi Jurgenson. 

1955. Putorius putorius orientalis. Polushina. Ekologiya, rasprostr., 
i nar.-khozyaistvenn. znachenie sem. kun'ikh zap.-oblastei 
Ukrainsk SSR, p. 6. 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. Nomen praeoccupatum — Mustela erminea 
orientalis 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 vis- 
ible. Lower side of body entirely blackish, without light under- 
fur; large yellow field absent on belly. Tail black throughout whole 
length. Postorbital constriction weak, its width in narrowest place 
not less or hardly less than interorbital constriction (V.H.). 

Description 

The general appearance, habits, etc. of the black polecat are typi- 
cal of other species of the genus. It also moves mainly in leaps, 
arching its back high upward, or, slinking down, it creeps. How- 
ever, 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 
hairs particularly long 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 8,500- 
9,000; ratio, one guard hair to 19-20 underhairs, and on abdomen 
average number of hairs per 1 cm^ about 6,000. Length of guide 
hairs on back about 50 mm, and thickness about 117 mk*; guard 
hair, correspondingly, 43 mm and 112 mk; and underhairs 26 mm 
and 20 mk (Tserevitinov, 1958). Winter fur of polecat is charac- 
terized by very great difference between length of guard hairs and 
of underhairs, which is not characteristic for the majority of spe- 
cies in the genus (except steppe polecat), and this causes guard 

*Microns. 



1109 

hairs to appear to protrude above underhairs. This is further accen- 
tuated by the contrast in color of the dark guard hairs and the light 
underhairs (see below). The tail at 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 is quite inten- 

sive brownish-black or blackish-brown, 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 two- 
layered color of the fur. The light undercoat is not seen equally in 
different parts of the body. On the back, especially the posterior 
half, long and numerous guard hairs completely or almost com- 
pletely hide the underhairs, and the light tone here is not notice- 
able or lightening is slight. On the sides, it [lightening] is well defined 
and their general color is strongly differentiated from the general 
tone of the [dark] spine. Approximately the same relation occurs 
on the neck. The nape and the shoulder region are darker in color. 




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 — black or blackish-brown throughout the entire 
extent. Sometimes, large blackish-brown fields are situated on the 
chest and inguinal regions and the belly 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 
black band. Ventral coloration characteristic of steppe polecat is, 
however, never observed (see [below]). Limbs are pure black or 
with brownish tint, the tail is black or brownish-black throughout 
its whole length. Light underfur is absent on the limbs and tail. 

A contrasting pattern occurs on the head — the area around the 
eyes, region between the eyes (anterior part of forehead) and lon- 
gitudinal stripe along the top of the nose 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. Vibris- 
sae are black. Hairs covering the lower side of feet are brownish- 
black 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- 
der the color tone of guard hairs and the lighter the general tone 
739 of the fur. The color of adult females shows no essential 
differences, but, apparently, their underfur does not carry rusty 
highlights and is 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 brownish- 
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 



fur 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 
the facial pattern is very variable — in some individuals, the head 
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 
it in the first days after leaving the nest. 

This pelage is converted in the first summer into the pelage of 
subadult animals. The fur is still somewhat "down-like", but guard 
hairs are well differentiated from underfur, the head pattern is "well 
defined, and contrast between color of underfur and that of guard 
hairs is clear. In general variation in color in this pelage is very 
great. It passes into the first winter pelage which does not essen- 
tially differ from the corresponding adult pelage (Herter, 1959,* 
Kratochvil, 1962 with modifications). 

Among exotypic variations of polecat are known complete al- 
binos and in some places, the chromic deviation known as 
"braginskii polecat" to furriers, and described as "P. p. stant- 
schinskii," is not uncommon. These chromists vary quite greatly in 
color details. In typical individuals, underfur is usually reddish, 
brighter than in normal polecats. Guard hairs on the whole trunk 
are bright and lustrous, reddish or brownish-red, or intensely red. 
The tail is reddish or reddish-brown. Black guard hairs are not 
everywhere, being absent on the lower body and head. The venter 
is light, having the undercoat color and only a large chest spot; 
inguinal region and posterior part of abdomen are reddish-brown. 
In extreme cases, guard hairs are so light that in tone, they are only 
slightly differentiated from the pale-yellow underfur or almost 
indistinguishable. In this case, the whole animal is 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 a sort of transition between normal 
individuals and chromists. 

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 
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 in adult 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 up- 
per skull profile in facial portion descends anteriorly as a convex 
arch, and the line of the braincase of skull gradually and slightly 
741 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 
very slightly diverging lines; narrowest place of postorbital portion 
not in form of sharp constriction — its width here equals to or only 
slightly less than interorbital width. Greatest constriction of post- 
orbital region with jaws closed usually lies posterior to line uniting 
apices of coronoid processes of mandibles."*^ 

Nasal opening is somewhat compressed laterally and width 
usually less than height. Auditory bullae somewhat swollen in pos- 
terior half; carotid 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 wid- 
ened 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 
throughout whole length of latter (noticeable only in skulls of young 
animals). 

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 
dimensions. In large individuals, all 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 protube- 
rances, crests, etc., not so wide and angular, and more narrowly 
separated zygomatic arches. Supraorbital processes usually smaller, 



■"According to Stroganov (1962), this is one of the most constant features dif- 
ferentiating it 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. 

In young animals (less than one year), cranial region of skull 
relatively much larger, convex and does not bear crests; supraorbital 
processes weakly developed, postorbital constriction weakly de- 
fined 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 
animals are well distinguished. 

Geographic differences in skull are not manifested. 

The OS penis, in general features, has a 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 
(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 differ- 
ence in dimensions of males and females, and the comparatively 
742 slow maturation of the young. They attain complete growth, appar- 
ently, only in the third year of life. Equally with this, is magnitude 
of individual variation in both males, and also females — but espe- 
cially the former. Besides these "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 polecats. Measurements of the latter ("giants") 
are considerably larger than those of polecats of the "normal" type, 
and lie 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 to very small individuals. The forms of these 
three types are also known in several other species of the genus, 
though in the 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 
been remarked on for "giants." 

Body length of males is 350-460 (up to 480 ?) mm, of 
females, 290-394 mm; tail length of males is 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 35- 
42 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 
is 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 
(17), 31.0-M34.0-37.0 mm; weight of penis bone is 0.280-M0.337- 
0.470 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.). 

Systematic Position 

The steppe polecat, M. (P.) eversmanni is the closest form to the 
black polecat. Their closeness is undoubted and is generally so 
great that a series of authors, beginning with Pocock (1936) in- 
clude them in one species. However, there is insufficient basis or 
principle for this and, evaluating all their particularities and all 
sides of their existence, it is 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 USSR itself represents a phenomenon whose nature is of a different kind. 

"■'After 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 
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). From 
this, the 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 mink is suffi- 
ciently close, although at a lesser level. The European mink itself 
represents, in a certain sense, a 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) to include minks in the genus of 
polecats (Putorius) as a separate subgenus. 
743 In West and Central Europe, a kind of domestic animal for the 

destruction of rats in houses and for hunting rabbits is kept, which 
is called the white polecat, Mustela furo.** This form is a typical 
polecat in its 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 skull of "M. furo" is generally typical of the forest pole- 
cat; however, in several cases, it bears features characteristic of 
the steppe (constriction of postorbital region). It has been noted 
that hybrids of black polecats 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 its 

♦Alternate name for steppe polecat, but also applied to domestic ferret (see 
below) — Sci. Ed. 

**ln English, "ferret" — Sci. Ed. 



1117 

morphology. "М. 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. p. furo L., Western Rif; accord- 
ing to Cabrera, 1932; Allen, 1939), This, apparently took place 
long ago, in a much earlier epoch, when it hardly seemed possible 
that 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.). 

Geographic Distribution 

Forest, forest- steppe, and in part steppe zones of the major parts 
of Europe, and extreme northwestern Africa. 

Geographic Range in the Soviet Union 

This itself represents the eastern half of the species range. 

The western border of the range of the polecat in our country 
constitutes the state frontier, from the mouth of the Donau [Dan- 
ube] in the south approximately to 62°40' N. lat. (northwest of 
Suoyarva), on the border with Finland in the north. 

In Karelia, the northern border extends from this point de- 
scribed above somewhat towards the southeast at first, approxi- 
mately to the Spassk Bay of Lake Onega, thereby passing around 
the West Karelian uplands from the south and then, passing around 
these uplands from the east, it suddenly ascends directly to the 
north passing in particular, near the western shore of Segozer and 
reaches Rugozer at 64*'10' N. lat. 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 it (Marvin, 1959; Parovshchikov, 1959). Since the 
polecat is spreading northward, it is possible that its distribution 
in eastern Karelia after the fifties and the beginning of the sixties 
has changed and the border extends farther north than is 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 
seacoast and here — almost at the Arctic Circle — attains the other most 
northerly point inhabited 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 

745 (near 63° N. lat.; V.Ya. Parovshchikov). Farther, the border appar- 
ently 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. 

The eastern border is very poorly known. Apparently, it ex- 
tends along the Urals, perhaps even to its eastern slopes, embrac- 
ing Sverdlovsk from the west. Old information on its occurrence in 
the Irbit region require confirmation. 

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 Тага 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 Magnitogorsk), 
this species, apparently, does not exist and here only the white 
polecat is found everywhere (Kirikov, 1952). Information concern- 
ing the occurrence of the described species along the eastern slope 
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» ПП ^ 



744 Fig. 265. Borders of the ranges of polecats in the European part of the USSR. 

V.G. Heptner. 
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 
Cis-Caucasus. Question marks in the Trans-Urals, — Irbit, in Kareliya — area of 
confirmed settlement in 50's and beginning of 60's. 



1120 

the shore of the Azov Sea, and along it, goes to the mouth of the 
Don. The black polecat is absent in the Crimea. 

In the steppes of the European part of the USSR, the black 
polecat is 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 
farther to the south, the stronger this association. The steppe pole- 
cat lives in the open steppe, and in forest very rarely occurs 
(Askaniya-Nova). 

From the mouth and lower Don, the range passes into the 
steppe of the western and middle Cis-Caucasus. 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 east- 
ern, a slightly curved line directed from Stavropol northwards to 
Tsimlyansk district on the Don'*^. This latter quite closely 
coincides with the borders of the Cis-Caucasian steppes. 

Details of the border of the range between the Don and Volga 
are not fully clear. Apparently, it suddenly begins to ascend north- 
ward along the Don, and then passes to the Volga somewhat south 
of Saratov. 

In the Saratov steppes of the Trans-Volga, the polecat is 
absent — it is encountered only in the extreme lower B[olshaya] 
and M[elenkaya] Irgiz. Farther, the border goes to the north along 
the Volga. Somewhat south of the Samara bend, it steeply returns 
to the east and, passing around Obshchii Syrt approximately along 
53** N. lat., reaches the Urals at the latitude of Magnitogorsk.'*^ 



■"The Cis-Caucasian part of the range is outlined here according to the individual 
occurrence on the map of Vereshchagin (1956). These points, in turn, were based on 
data 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 sometimes still denied 
(Kuznetsov, 1952). The possibility is not excluded that this species may penetrate as 
individual animals 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 to the south of the line Zhitomir- 

Kiev-Orel-Voronezh-Gor'kii-Kirov-Ufa-southern Urals (approxi- 

746 mately), i.e. in its greatest part, overlaps the range of, the steppe 

polecat. In this region, individual hybrids are encountered (quite 

rarely), but hybrid populations are not formed anywhere. 

The black polecat itself represents a species which is notice- 
ably widening its range towards the north. Its 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. However, there are no accurate data 
concerning its movement in this direction. 

The tempo of colonization to the north is quite energetic. 
Thus, in Karelia, from 1930-1932 to 1956, the polecat colonized 
towards the northwest about 250 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 Severnaya Dvina from the 
latitude of Shenkursk to Arkhangelsk, i.e. for a distance of about 
320 km in a straight line (Parovshchikov, 1959). The polecat 
appeared at Kaisk Pochinki on the Kama at 60° N. lat. at the end 
of the 20's of this century (Lobachev, 1930) and by 1956, it 
began to be found, though not often, in the upper Vychegda 
(Parovshchikov, 1959). Therefore, it moved about 300 km north- 
ward or northeastward during a period of approximately 22-25 
years. Within 25 years, it moved 300 km along the Onega. Such 
a tempo of colonization — on average about 10-12 km per year — 
is considered to be very high. 

Colonization is mainly stimulated, apparently, by felling of for- 
ests and 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- 
der in several segments, a more or less significant "oscillations" of 
the border occur locally. 

Geographic Range outside the Soviet Union 

This occupies almost all of Europe. In the north, the border in- 
cludes Finland south of 66° N. lat.; in the Scandinavian Peninsula, 
it begins at the Baltic coast at almost 60° N. lat. descends a little 
westwards including Lake Vattern* from the south, in southeast- 
ern Norway, it forms a narrow projection towards the north ap- 
proximately 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 

*In Russian original, "Venern" — Sci. Ed. 



1123 



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, 
the Dobruja region (V.H.). 



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Г959 




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747 Fig. 267. Movement of the northern border of the range of the black polecat in 

Arkhangel'sk district (Parovshchikov, 1959, with additions). Lines designate the 

general limits of the range-points — the individual appearance of animals far from the 

general border. The movement of the border does not fully correspond to that which 

was established for Karelia. V.G. Heptner. 



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. It is possible that the black 
polecats of the steppe zone are somewhat lighter than those of the 
forest zones (Brauner, 1929). However, these differences apply 
only to a very small part of the species population in the USSR; 
they are not yet proved sufficiently and are so insignificant that the 
southern population does not deserve special designation. If this 
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 Euro- 
pean 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). A 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 in the European 
part of the USSR, some geographical localization of the red mu- 
tant "P. stantschinskir is recognized. Individuals of this form are 
known from Smolensk district (in particular, Roslavl' region), 
Tambovsk, Moscow district (Klin region), 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 
(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 approxi- 
mately 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. (P.) p. putorius Linnaeus, 1758 
(syn. — vulgaris, foetidus). 

Body and skull dimensions larger than Middle Russian form, 
fur darker, lustrous and fluffy. 



1126 

Found in western part of European territory of USSR to the 
west from the given line, i.e. Pribaltika; Western Byelorussia, 
western Ukraine. 

Outside the USSR, found in central 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 
Russian polecat are associated with its relatively recent coloniza- 
tion of the territory it now occupies (colonization beyond the Ural 
is, apparently, continuing). 



Outside the USSR, the following forms are usually recognized: 
1) 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 — Dobruja, Romania; 
4) M. (P.) p. angliae Pocock, 1936— England; 5) 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 its 
distribution within the range, these may be judged by indirect data — 
the figures of commercial catch. Before the October revolution, 
about 150 thousand skins of 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 
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, Moscow, 
750 and Ivanovsk districts and from the Ukraine. Indices of catch from 



1127 

these ranged (in 10 km^) from 0.028 in former Severnaya Terri- 
tory 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 great- 
est density within the boundaries of the USSR is observed in the 
western part of the range, which is explained by the favorable 
habitat conditions for this animal there. 

Habitat. The forest polecat avoids large, continuous forest 
massifs. Small forest islands and isolated groves alternating with 
meadows, cultivated fields and human settlements are more suit- 
able for its life. It is 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 forest creeks, and they are 
generally restricted to relatively populated regions. Together with 
human cultivation and agriculture, it gradually penetrates into the 
taiga. 

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, boggy 
depressions. Without penetrating deep into the forest, it adheres to 
openings, borders scrub forest and old clear-cuts, especially those 
adjacent to worked fields and settlements. Rarely it may be met 
with in mossy bogs or in deep forests. It settles 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- 
stricted to settlements, isolated farms, 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) 
vole [Microtus arvalis] occupies first place (18.5-26.8% occur- 
rence). Red-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, espe- 
cially in winter, are amphibians — mainly, grass frog and green 
toad. 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): domes- 
tic hens and pigeons, quail, gray partridge, grouse and various 
small birds. The role of the forest polecat in the destruction of 
poultry is greatly overestimated. Special investigations showed that 
in kolkhozes [cooperative farms] and poultry sovkhozes [state 
farms], losses from it ranges from 0.2 to 2.5% for the adult birds 
and from 0.5 to 12%, for pullets, most often in the haying period. 

Among a number of rare foods of the polecat are the hedge- 
hog, adder [Vipera aspis], grass snake [Tropidonotus natrix] and 
insects. The significance of individual foods changes depending 
on fluctuation in 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 car- 
rion, 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 fish (0.9%) (Grigor'ev and Teplov, 1939). 



751 Table 68. Geographic variation in foods of the forest polecat (in percentage of 

occurrence) 



Type of food 


Volga-Kama Territory (Gi 


rigor' ev 


Middle zone 




and Teplov, 1939) 


1 


(Lavrov, 1935) 




Summer 


Wintei 




Winter 




M 


lim 


M 


M 


Mouse-like rodents 


28.8 


42.0—65.0 


25.4 


51.2 


Water vole 


89.1 


3.7—32.1 


14.7 


2.4 


Hamsters and ground squirrels 


— 


3.3—9.8 


— 


— 


Insectivores 


— 


0—8.0 


4.1 


2.4 


Birds 


— 


0—12.2 


8.0 


16.3 


Amphibians 


— 


0—20.0 


12.1 


19.5 


Fish 


0.9 


0—16.0 


6.9 


0.8 


Carrion 


— 


— 


— 


9.8 


Number of records 


114 


563 


— 


123 



1129 

In the western districts of the U[krainian]SSR (analysis of 1 11 
stomachs and 31 feces), mouse-like rodents predominated in the 
food of the forest polecat, though the occurrence of all 18 com- 
ponents was not higher than 10% (Polushina, 1958). 





% 




% 


Hedgehog 


1.8 


Gray vole 


7.6 


Shrew 


2.6 


Domestic rabbit 


7.6 


Norway rat 


6.8 


Sparrows 


7.6 


House mouse 


7.7 


Song birds 


5.0 


Harvest mouse 


0.9 


Bird eggs 


3.4 


Red-backed vole 


3.4 


Amphibians 


9.3 


Mice, Apodemus 


6.0 


Fish 


4.9 


Water vole 


3.4 


Carrion 


4.2 


Root vole 


1.8 


Insects 


5.0 



The forest polecat characteristically makes food stores. The 
bodies of water where grass frogs overwinter are used as "store- 
houses" by polecat in the winter time. The polecat continuously 
visits such places. 

Home range. In contrast to the steppe polecat, the forest pole- 
cat lead a more settled way of life and has a definite individual 
home range. However, the dimensions of the range are unknown 
nor is there data on length of its 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 5 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 is an animal 
with a crepuscular and nocturnal rhythm of daily activity. It is very 
rarely active during daytime, only when motivated by hunger. This 
is confirmed by laboratory experiments (Kalabukhov, 1943): the 



изо 

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 
months) differ greatly: it is equally active throughout the course of 
752 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 
than solongi, ermine and weasel — man in condition can catch a 
running polecat. The sense organs are well developed but it cannot 
distinguish between different colors. 

Seasonal migrations and transgressions. Only small seasonal 
migrations are known but far from all polecats participate. In au- 
tumn, many polecats move to villages, attracted by the concentra- 
tion here of mouse-like rodents after the gathering in of the harvest 
from the fields. In spring (April), a reverse movement is observed. 

Reproduction. As in the steppe polecat, the period of sexual 
activity extends from the second half of February to the second 
half of June. Evidently, the rut often extends into April-May 
(Lavrov, 1935). Duration of pregnancy is 42 days (Moscow Zoo; 
Manteifel', 1947). 

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 
sparse, short whitish down. Eyes open on the 34th-36th day; at 
that time, the wool darkens. Sometimes, the brood does not dis- 
perse until spring. Sexual maturity is attained at about one year, 
but full development and dimensions at two years. Molt in the 
forest polecat occurs twice annually — in spring and autumn. 

Enemies, diseases, parasites, mortality, and competitors. Com- 
petitors 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 
occur. Bodies of polecats, dying from causes that are usually un- 
clear, are more often found than other carnivores. The forest pole- 
cat is susceptible to canine distemper, and it suffers from an 
unknown infectious intestinal disease accompanied by high mor- 
tality. A number of helminths were recorded, including not less 
than four forms of nematodes parasitizing the frontal sinuses and 
nasal cavity. In France, distomatosis was revealed in every fourth 
animal. In some cases, the infection reaches the brain and causes 
the death of the animal. Usually, many fleas and ixodid mites 
occur on the forest polecat. In captivity, the forest polecat lives up 
to 12 years (ManteifeF, 1947). 

Population dynamics. The forest polecat is characterized by 
significant fluctuations in numbers. Their cause and character are 
unstudied. From 1928/29 to 1949/50, i.e. for 22 years, numbers of 
forest polecat in the southwestern part of Kalinin district increased 
three times, but to different levels. The increase in the number in 
1945/46 was three times smaller than the rise in 1929/30 and six 
times smaller than the maximum rise of 1938/39. 

There is no strict periodicity in the population fluctuations. 
Apparently, the period is about 7-9 years. Reduction in the popu- 
lation 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 in the period 
1939/40 to 1940/41 fell 9-fold. 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 cm and prints of claws and 
callosities of the feet are very distinct (Formozov, 1952). For dif- 
ferences from tracks of mink, see page 1106. Triple prints of the 
feet are as typical for the forest polecat as pairs. When walking, it 
frequently changes feet, and often one pace for another. The gen- 
eral character of the track or trail is distinguished by an absence 



1132 




753 Fig. 269. Tracks of the hind and fore feet of black polecat on silt. Shore of Moscow 
river, Zvenigorod. 22 June 1939. Sketch by A.N. Formozov, about 2/3 natural size. 

of elaboration typical of the mink. It moves in energetic leaps, digs 
in places inhabited by mouse-like rodents, and prefers to catch 
animals moving on the surface, so rarely digs in the snow. Its track 
is not so tangled as weasel and ermine (Formozov, 1952) (P.Yu.), 



Practical Significance 

The forest polecat is a valuable fur-bearing animal. The price of its 
skins is higher than that of steppe polecat. The fur enjoys a good 
demand on the world fur market on which, before the 1914-1918 
war, Russia offered more than 50% of all skins. After the October 
revolution, the take of this species in the whole USSR increased 
noticeably as reckoned by growth of the catch, and also as calcu- 
lated by a decrease in its number in western European countries. 
In the years after the 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 1926- 
1929, 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 
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 commercial hunting, and for the most 
part incidentally captured. 

It is necessary to restrict the hunting of the forest polecat to 
the period of complete "primeness" of its 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, p. 
447. Utsang in southern Tibet, north of Sikkim. 

1910. Putorius eversmanni Lesson var. michnoi. Kastschenko. 
Kashchenko. Ezhegodn. Zool. muzeya Ak. Nauk, 15, p. 271. 
Kiran river 20 km from Troitskosavsk (Kyakhta), Trans- 
Baikaliya. 
754 1912. Putorius eversmanni sibiricus Kastschenko. Kashchenko. 
Ezhegodn. Zoolog. muzeya Ak. Nauk, 17, p. 395. Aginsk 
steppe, Trans-Baikaliya. Nomen nudum. Nee Mustela 
putorius sibiricus Radde (= Mustela sibirica Pallas). Name 
given with authorship of Radde (1862)^°. 

'"Kashchenko' s uses of the cited name is evidently a misunderstanding. 
Radde (1862, 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 diag- 
nosis. Apparently, he had a winter individual in one case, and in the other — 
a summer, or an unmolted one. 



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 i 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. 

11, p. 25. Blagoveshchensk on Amur region. 
1936. Putorius putorius aureus. Pocock. Proc. Zool. Soc. London, 

p. 703, Kazan.^' 
1958. Putorius eversmanni nobilis. Stroganov. Izv. Sibirsk. otd. 

AN SSSR, 11, p. 150. Vic. of Kokchetav, Kazakhstan. 
1958. Putorius eversmanni pallidus. Stroganov. Ibidem, p. 150. 

Kargat, between Novosibirsk and Lake Ghana. 
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. 

inst. Sibirsk. otd. AN SSSR, 6.** Hi river, southern 

Pribalkhash'e (V.H.). 

Diagnosis 

General color light, whitish-yellow, only weakly overlaid with 
blackish-brown (dark guard hairs). 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, 
apparently, not accurately reported, or is incorrect — the individual was received by 
the British 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 
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 inguinal region 
brown in color. Base of tail light, only terminal part dark-brown. 
Postorbital constriction of skull well marked — narrower than 
interorbital (V.H.). 



755 



Description 



The white polecat is entirely similar to the black in general appear- 
ance, proportions and habits, but its body appears somewhat more 
elongated. Possibly, this is due to guard hairs, especially those 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 
length, covered by short close-lying hairs, and on the whole, is 
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, is usually less in steppe polecat 
than forest polecat. Also less is general thickness of pelage layer. 
On the whole, fur of steppe polecat is somewhat coarser than for- 
est polecat. Guard hairs more strongly developed and denser in 
posterior portion of dorsal 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, their secretion has same sharp 
smell as in black polecat and, apparently, also can be sprayed (no 
direct information on this). 

Teats, 3-5, usually 4 pairs. 

General basic color tone of winter fur very light, yellowish or 
whitish-yellowish, determined by underfur color. Over this 
background, is 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 
the back, 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 re- 
gion, 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 
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. Mask 
often narrow and pale, dark region is light and whole head light. 
Sometimes, head is entirely white or almost white (slightly ocherous) 
and head pattern is absent. 



1137 

Lateral parts of neck of same color as anterior part of sides 
or lighter; its lower surface, posterior to throat dark, blackish- 
brown or brown. Chest and forelegs black or blackish-brown; 
Venter light, yellowish-straw; its posteriormost part, inguinal area 
and posterior extremities dark, black-brown — as in fore limbs. 
Basal part of its tail (half or two-thirds) has light color tone like 
that of sides; distal part brown or black-brown, almost black. 

Vibrissae black and white; hairs covering soles of feet close- 
fitting and elastic, black-brown and hiding digital callosities and 
foot pads. Claws light-horn [in color]. 

Fur color is 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 darken- 
ing 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 is a 
median line on the venter, a narrow dark stripe extending between 
dark areas on chest and groin. 

Summer coat shorter and coarser than winter, and is not as 
dense and close-fitting. Ocherous or reddish tone strongly devel- 
oped. 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 pie- 
bald. 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. 

Newborn polecats are naked, but in first 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 development begins 
earlier and proceeds more quickly than in the forest polecat, and 
it is already fully developed in the subadult individuals (Kratochvil, 
1962). In polecats of intermediate age, mask and entire head pat- 
tern are well-manifested — they are relatively clearly outlined and 
densely colored. With age, the whole head lightens, parietal and 
occipital parts disappear or become paler, their area decreases and 
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 
and neck strongly lighten. With age, general tone of underfur light- 
ens, which may lead to almost white [pelage] with light straw- 
colored frosting. 

"Exotypic" variations which are so sharp in black polecat are 
not noted in the steppe, or they are rare. In particular mutations 
like the "braginsk" polecat have not been described. 

Geographic variation in color, in contrast to that observed in 
black polecat, is better manifested, and reveals itself in degree of 
757 intensity of general tone of underfur (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 parallelisms of not only individual and age variation, but 
also geographic variation is manifested. Density and length (qual- 
ity) 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. Mas- 
toid 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 sepa- 
rated 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 its postorbital part 
not parallel, but form two lines which meet at an angle in a sharp 
constriction (interception; "waist") and behind it, again diverge. 
Width of this part in its narrowest place is less than interorbital 
width — in old individuals considerably less. Most constricted part 
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 all canine and carnassial teeth, 
generally more massive. In connection with this, lower jaw also 
somewhat more massive. 

Female skull differs from skull of male (not to mention meas- 
urements) 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 ante- 
rior, closer to supraorbital processes. Braincase more swollen and 
narrower posteriorly, without crests or only just noticeable. In 
connection with relatively wide interzygomatic area, whole brain- 
case 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 — the skull of the subadult steppe polecat is similar 
to that of a more mature black polecat in some respects (chiefly, 
structure of interzygomatic area). Further, in connection with con- 
striction of postorbital 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 more rapid. In one way or another, the con- 
trast in skull structure of young and extremely old steppe polecats 
(with almost obliterated teeth) is much greater than that in black 
polecats of their respective ages. 

As a very rare individual structural deviation, there is in the 
skull sometimes a bony connection between the end of the ptery- 
goidal 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, in general, greater than in the black 
polecat. This is, apparently, associated not only with the longer 
growth period in the steppe polecat, but also with some geographic 
variation in this character, 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 exploi- 
tation is practically undeveloped, and the animals live to the end 
of life, or in all events, grow to their maximum limit. The black 
polecat is hunted quite intensively everywhere. 

As in black polecats, giant individuals are encountered among 
white polecats, the measurements of which extend beyond the lim- 
its of "normal" fluctuating variation in the species. They are met 
with significantly more often, at least in some places, than the 
black polecat. They are most of all found in 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 tail end and brown feet; 
I.M. Zalesskii, 1930), it is 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. 

^^In the steppes along the upper Irtysh, they are even known to the local inhab- 
itants who call the abnormally large polecats "mogil'shchik" (=grave digger) because 
of the belief that they live in graveyards and feed on the dead (Zverev, 1931). 



1142 

Sexual dimorphism in dimensions is quite sharp — female body 
length consitutes about 85-90% of male, and weight averages about 
45-50% of male weight. 

Body length of males is 320-562 mm, of females, 290-520 
mm; tail length of males is 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 
(Stroganov, 1962). 

Condylobasal length of male skull is 61.7-82.2 mm, of 
females, 52.4-76.7 mm; zygomatic width of males is 30.0-58.9 
mm, of females, 30.0-48.7 mm; mastoid width of males is 35.7- 
47.9 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 transgres- 
sion in length, no transgression in weight of both age groups is 
observed). 

Weight of Siberian males up to 2,050 gm, of females, 1,350 
gm (Stroganov, 1962). 
760 One giant polecat had a body length without tail of about 
75-80 cm, although normal dimensions of animals from this re- 
gion (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 at their disposal, genally speaking, only insignificant mate- 
rial of the white polecat. The idea was not met sympathetically by 
us. Actually, if all features and properties of both polecats are 



1143 

evaluated, and the question is approached not only from a purely 
morphological point of view, it 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 argument 
favor the treatment employed here — M. (P.) putorius 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 dentition, 
protuberances, crests, etc. are better developed and the masticatory 
("predatory") musculature is 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 indi- 
vidual races, is larger than the black polecat. It is naturally as- 
sumed that all of this is related to larger prey. The close connection 
of this polecat with ground squirrels and apparently even marmots 
in some places is better recognized and understood. It is associated 
with murid rodents to a lesser extent than the black polecat and, 
apparently, is completely independent of amphibians. 

The course of age changes in both species shows that their 
skulls are very similar in early age — no essential differences exist 
between them in the interzygomatic region. Later, in the black 
polecat, the above-described structure of the interzygomatic region 
is formed, and at this age changes in this region terminate. As 
mentioned, these changes go farther in the steppe polecat, with 
significant changes appearing, all 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 men- 
tion females and males of younger ages at the same chronological 
stage as white polecats. The correlation is approximately as in 
European and American mink, but the differences in morphology 
of the final stages are greater. 

Geographic features of both species obviously give evidence 
for their treatment as independent species. However, if the entire 
range of both species is taken into account, a definite geographical 
761 vicariance is delineated, although the region of sympatry of the 
two species is great. Together with this, the boundary "transitional" 
belt is not narrow, but a significant part — about one-third if not 
more of the range of the black polecat — is covered by the range 
of the white (the reverse correlation is different). Both species 
exist in this region of mixing, in a known degree of independence, 
inasmuch as there is the general possibility for mammal species 
with this type of ecological isolation. The steppe polecat confines 
itself to open expanses, and the forest — closed biotopes or human 
settlements. In the southern European part of the USSR, the black 
polecat penetrates into the steppe zone chiefly along the valleys of 
rivers, and in the steppes, with ground squirrels, the white polecat 
lives. Thus, along the Dnestr [river], in its 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 — the 
black, and to the south in the Carpathian plain — the white 
(Konyukhovich, 1953), etc. Both species exist together in Czecho- 
slovakia and Austria, but in different biotopes (the white — in steppe- 
type biotopes with ground squirrels) (Bauer, 1960; Kratochvil, 
1962). They also colonized the Russian plain in the east and north 
(see later); the white polecat, as said, freely penetrated into 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 — forms the species independence relative to the 
polecat is 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, in their entire territory within the limits of 
the USSR (apparently, and outside the USSR) black and white 
polecats are encountered together, and nowhere do they produce 
hybrid populations, and a zone of transgression between the two 
species is 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 ferret* (M. (P.) nigripes Audub. et 
Bachon.) is 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 
dimensions of the separate parts. 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 
the chest region, lower part of neck and inguinal region. It is 
entirely likely, that if the 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.). 

762 Geographic Distribution 

Found in Central and in part in Middle Asia, Kazakhstan, southern 
Siberia; southern and in part middle parts of eastern Europe and 
Central Europe. 

'■•It is possible that 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). 

"For the systematic interrelationships of polecats and related species of Eurasia, 
see also the corresponding sections on black polecat, 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 
the species, 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; 
Konyukhovich, 1953). Apparently, it occupies the entire Trans- 
Carpathian plain, but is absent in the mountains. This area of 
occupation of the white polecat is isolated from the remaining 
range of the species within the boundaries of our country and is 
connected with its central European part lying in Czechoslovakia, 
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, evi- 
dently, 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 north- 
ern 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, direct- 
ing itself towards Bryansk and even somewhat northwest of it — to 
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, 
Ryazan' and Gork'ii (Kuznetsov, 1952); i.e., approximately along 
the Oka [river]. According older data, the occurrence of the steppe 
polecat was recorded south of the line leading eastward from Orel, 
at Chaplygin (Ranenburg), northwest of Tambov, at Morshansk 
north of 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 every- 
where, even the most northern regions of the district and was not 
recorded only in the very northwestern bordering Moscow, Yaroslav 
and Ivanov districts — in Aleksandrov — and in one of the south- 
western [regions] — Kurlovsk (in Vladimirsk Meshcher to the west 
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 it is 
not exclusively a rarity in Vladimirsk (see later). 

It is possible that the light [white] polecat penetrated into the 
southern parts of Gork'ii [in] Zavolzh'e and exists in this district 
764 not only south of the Volga. In any event it was recorded not only 
for Chuvash, but also in the eastern part of Mari ASSR (Yurgenson, 
1932). Therefore, reaching the Volga somewhere above the mouth 
of Kama, the borderline directed itself quite steeply to the north- 
east, covering not only the entire Tatarsk Republic, but also the 
southern part of Kirov district. In an unclear way it is directed 
closer to Omutninsk (5840' N. lat. and a little east of 52° E. long.; 
Popov and Lukin, 1949). This is the northernmost point inhabited 
by the species in the European part of the country. 



*In Russian original, misspelled "Telyache" — Sci. Ed. 

'*Due to technical reason, the maps pubUshed by Heptner et al. (1950, 1956) 
contained an error — it contained no data on the occurrence of the white polecat in 
Moscow district and is absent in Smolensk. In agreement with the new data, the 
border in the east extends in a different way. 

In 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 the east of the European part of the USSR — in Priural'e 
and in the Urals, the border of the range is very poorly clarified. 
From Omutninsk, it apparently is directed towards the Urals (the 
white polecat does not reach the northeastern parts of Perm 
district; Kuklin, 1951) and along its western foothills, and encom- 
passing 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 east and crosses the Urals. The assumption that the northern 
border of the range passes through the Urals below 58° N. lat. 
(Yurgenson, 1932), i.e. that the steppe polecat lives all over the 
middle and southern Urals, is apparently wrong. 

In the Trans-Urals 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 a point a little north of it. But, its occurrence so far 
to the north in Trans-Urals sometimes gives rise to doubts 
(Stroganov, 1962), and it has been assumed that the border of the 
range from Sverdlovsk goes to Irbit and thence even to Tobol'sk. In 
any event, in Trans-Urals the polecat does not reach Ivdel' (60° 40' 
N. lat.). 

From Tobol'sk, the border line, generally following the Irtysh, 
or going a little to its north, directs itself east approximately to the 
mouth of the Shish river and there it turns to the sources of 
Vasyugan (I. Laptev, 1958), including, in this way, the entire basin 
of the Тага (Yakushevich and Blagoveshchenskii, 1952). From the 
source of Vasyugan, it again extends to the east or, somewhat 
deviating to the north, and reaches the Ob' (the polecat is absent 
to the north of the 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, 
the polecat was recorded at Narym (Ognev, 1931) and in 1947, 
one polecat was taken on the Ob' at Lokosov (Lokusov), some- 
what 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." 

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 it from the north. For the Yenisei 
basin, there is evidence of the capture of a polecat at the conflu- 
765 ence of the Viv' river with the Nizhni [Lower] Tungusk (at 64°; 
Yakovlev, 1930; Ognev, 1931); this is a poorly understood mis- 
take — the range undoubtedly does not extend so far. 

To the east of Baikal, the range on the north includes the 
region of the left tributaries 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, S. 
Ananshkin). These are peripheral points where the polecat is very 
rare and is 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 km 
along the river. The polecat occupies a similar, if not smaller, area 
along the upper Angara near the mouth of the Kuta (V.V. Timofeev). 

In the east, 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 at the [Siberian] railroad 
mainline). Thence, it steeply descends to the south to the Argun', 
nearly in the region of the mouth of the Gazimur, and exits beyond 



"Polecats were found at some points on the Ob' between the mouth of Тут 
(80° E. long.) and Surgut; i.e. north of 60° N. lat. were apparently, wrongly 
placed on Stroganov's map (1962) — these were not clarified in the text and 
contradict it. 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 
of Tomsk district. 



1151 

the frontier (Favorskii, 1936; P. Zimin). Therefore, it is evident 
that the polecat 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 separate small area inhabited by the polecat in our country 
is located on the middle Amur. It occupies Blagoveshchensk 
region 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, 
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 
the Trans-Baikal region through this country. References to its 
occurrence along the Uda (Radde, 1862; Ognev, 1931; Stroganov, 
1962) are apparently, mistaken. This error may be, at least par- 
tially, due to the fact that the data concerning the Uda — a tributary 
of the Selenga (Baikal basin) — where the Siberian polecat is com- 
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 Azov seas, and in the Crimea, the steppe polecat 
lives not only 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, approxi- 
mately 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. The pole- 
cat does not ascend the mountains (the foothills are wooded or 
were wooded) and is known to a height of only about 600 m 
(Psebai) or even less (Dinnik, 1914; Vereshchagin, 1947, 1959; 
V.G. Heptner). Therefore, in the Main Caucasus and in the Trans- 
Caucasus, the polecat is absent. In the regions adjacent to the 
northwest, north and northeast of the Caspian the range extends to 
the seacoast. 

Precise borders of the steppe polecat range in Middle Asia still 
766 cannot be established. In any event, it is far from being distributed 
everywhere here, and the line of the southern border of the range 
in this country is quite complicated. The polecat exists bx)th in 
Mangyshlak and on Buzach peninsula (Ognev, 1931; Gerbel'skii, 
1939; Sludskii, 1953) and along the Caspian Sea (Kara-Bogaz- 
Gol; M. Laptev, 1936) the range descends to Krasnovodsk as a 
narrow extension and, perhaps, may go along the Sea and a little 
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, ap- 
parently, along Ustyurt Chink in which the polecat exists, particu- 
larly in its 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 
south, the polecat is distributed along the rivers, apparently, to 
Khiva and Urgench (Gladkov and Nikol'skii, 1935; Kostin, 1959; 
Ishunin, 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 
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 is, appar- 
ently, absent. 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- 
ently, along the entire 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 south- 
ward 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; 
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 km to 
the northeast of Bukhara). Farther south, the polecat is known in 
the Karsha, Guzar (southeast of Karsha) and Saryassii (near Denau 
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). 

Based on these places of occurrence, the border of the polecat 
range in the Trans-Amu-Dar'ya part of Middle Asia can be, appar- 
ently, preliminarily be given thusly. From the lower Amu-Dar'ya, 
probably at the level of Khiva or somewhat below, the border line 
includes parts of the desert directly adjacent to the delta (in par- 
ticular, regions of old cultivation) and then turns northeast, con- 
tinuing along those parts directed towards the Aral Sea, and crossing 
the Dzhandar'ya and Kuvandar'ya districts, reaches the Syr-Dar'ya, 
probably, somewhere in the Kzyl-orda district. Being directed to- 
wards 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 
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, moving eastward, goes 
on to Surkhandar'ya. Thence, in a still completely unclear form, it 
767 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 
occupies the whole Kzylkum, which is, however, evidently incor- 
rect (T.Z. Zakhidov). 

Unique in the sense of its general outline, the range of the 
species and, to a considerable degree, its 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 is associated, probably, in addition to biotopic con- 
ditions, with some of our mountainous regions which the polecat 
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, how- 
ever, do not exist. 

The range of the white polecat has noticeably broadened west- 
wards 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 begin- 
ning 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 
western Ufimsk), though it is not excluded that he did not always 
differentiate 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 
the beginning of our century penetrated into Tatariya but did not 
still occupy it entirely. Near Kazan, it appeared at the end of the 
20' s, and this was the most northerly point of occurrence of the 
species known at that time (Ognev, 1931). In the Omutninsk re- 
gion of Kirov district the polecat apparently penetrated in the 40' s 
of our century. If these indications perhaps do not completely 
accurately describe the chronology of movement of this species to 
the west and north in the Trans- Volga, in any event it can be 
considered that the tempo of its colonization is 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 
excluded that its settling proceeds not only from the south but in 
part from the east. 

It 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 Europe is also 
associated with recent times: in 1928 it was established in Hun- 
gary (Ehik, 1928), after several years in Trans-Carpathia, in 1948 
in Czechoslovakia and in 1952 — in Austria where it rapidly spread 
and its number strongly increased (Bauer, 1952; Kratochvil, 1962; 
O. Wettstein)^^ 
768 It is difficult to establish the time 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 all of this, Pleistocene fossils of polecat (M. 
ev. soergeli) are known from France, Baden, Wiirttemberg, Hun- 
gary and Austria and the Pleistocene and Holocene from the south- 
ern European part of the USSR (Ekhik, 1928; Pidoplichko, 1951). 

'4n light of the stated facts, information on the occurrence of steppe polecat near 
Warsaw (apparently in the last century; Ognev, 1931) is 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. Only a short time ago, the white polecat had slightly 
penetrated into Poland from Rava-Russkaya region. 



1156 





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768 Fig. 273. Species range of the white polecat, Mustela (Putorius) eversmanni Less. 
Southern and eastern borders in 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 some reason abandon range. 

If it is now difficult to say anything about the time of the 
appearance of the polecat in the southeast and the extreme south, 
and in part in central Europe, then, in 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 of the range to the north is occurring locally 
also in western Siberia; however, the progressive movement of the 
border there is very slow. In Siberia, colonization of polecat in 
new places is, apparently, directly associated with the cutting down 
and clearing of the taiga. In Europe, this cause also has certain 
significance (Trans-Kama); however, in many places, colonization 
of the polecat in many places is, apparently, determined by other, 
apparently partially internal reasons. This is 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 distribution of the polecat in Cis-Baikaliya and 
Trans-Baikaliya among separate isolated small steppe areas (Lena, 
upper Angara and others) permits one to think, on the contrary, 
that the distribution of this species in Pribaikal was never large. 

For the sake of accelerating colonization in Siberia, some attempts 
were made to acclimatize the polecat somewhat more to the north of 
the border of its natural range (Narymsk district; Lavrov, 1946). 

Geographic Range outside the Soviet Union 

In Europe, it occupies Romania (apparently, except montane wooded 
parts of the Carpathians), Hungary, western (Lower) Austria, 
Czechoslovakia, northern Bulgaria and northeastern Yugoslavia 
(Serbia). 

In Asia, the range occupies Dzhungaria, the Mongolian Re- 
public, Kashmir 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, 
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 montane regions of Tibet. Gen- 
erally, the range in Asia outside the borders of the USSR is very 
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). 

A 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 irregu- 
lar 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 the vast range, geographic variation of the white 
polecat is, relatively, not as great as might have been expected, 
especially taking into consideration that the animal is met with in 



1159 

very different natural regions. It is possible, that part of this 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 dimen- 
sions (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 is, obviously, too great and 
gives an erroneous and exaggerated impression of geographic vari- 
ation of the species. Even the number of the recognized fur grades 
is considered to be too large (Kuznetsov, 1952). 

A full revision of geographic variation of the species is neces- 
sary. 

"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- 
scribed forms and may be completely distort them. Compare, for example, illustrations 
and descriptions of the forms lineiventer and tuvinicus. 



1160 

The enumeration below must be viewed as completely prelimi- 
nary, of designated forms, in part good and in part weak. Among 
them, three groups are more or less clearly recognized: 1) polecats 
of Europe, western Siberia and northern part of Kazakhstan — group 
hungarica-eversmanni; 2) polecats of Trans-Baikaliya and Pri- 
Baikaliya, Mongolian Republic, Altai, mountains of Middle Asia 
and Tibet — group michnoi-larvata; perhaps the less known pole- 
cats of the plains of Middle Asia and southern Kazakhstan 
771 {''talassicd") may be related to this group, but more probably they 
belong to the first group; 3) polecats of Priamur'e and the eastern 
part of former Manchuria — amurensis. 

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 dark- 
tawny tips of guard hairs. Neck grayish-yellow, tail half sandy 
color, half dark-brown. 

On the whole, color darker than in nominal form and other 
races. 

Information on dimensions absent. 

Westernmost part of range in European part of country, north- 
wards apparently, to northern limit of distribution, eastwards, prob- 
ably to Volga and northern Caucasus. 

Outside the USSR — in Czechoslovakia, Austria, Hungary, 
northeastern Yugoslavia, northern Bulgaria, Romania and appar- 
ently in a small section of Poland at its boundary with L'vov dis- 
trict. 

There has been no direct comparison of materials from south- 
ern Russian polecats, described as occidentalis, with those of Central 
Europe; however, their diagnoses are actually identical, and they 
must, apparently, be considered identical. The possibility is not 
excluded that the range of 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 mod- 
erate length. Length 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 
the basic light color tone or darken it a little. Neck whitish. Tail 
not heavily furred, its basal part mainly whitish, the distal third 
black-brown. 

Body length of males is 320-460 mm, females, 280-410; tail 
length of males is 80-160 mm, females, 70-140 mm; length of 
hind legs of males is 40-70 mm, females, 30-54 mm (after Zverev, 
1931 and Stroganov, 1962). 

Average dimensions (after Zverev, 1931): body length of males 
(561) 390 mm; females (671) 340 mm; tail length of males (519) 
120 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 12.0- 
16.6 mm, females, 11.3-15.2 mm (skull measurements after 
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 possibly be encountered that part of China contiguous with 
eastern Kazakhstan. 
772 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 — in Trans-Volzh'e polecats, for instance, the black tip 
occupies half the length of the tail, and the skin is quite darkened 



1162 

by the guard hairs. The limit of the range in the east is also poorly 
defined. 

The form "P. p. aureus" Pocock, 1936 described from near 
Kazan, is 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.) ev. talassica Ognev, 1928. 
Dimensions small. Winter fur long and dense, but somewhat 

coarse. General color very pale, whitish, without rusty tones; 
underfur slightly yellowish, guard hairs pitch-black, but length of 
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. 

A very little known form, apparently occupying the plains of 
Middle Asia and the southern belt of Kazakhstan, not penetrating 
high into the mountains. It is, possibly, related to the group of 
Central Asian races (michnoi — larvata; Ognev, 1931); however, it 
is evidently smaller and apparently is allied with those described 
above. 

In the fastness of the Pamiro-Alatisk system (Alaisk valley), 
and possibly deep in the Tien Shan another form exists (see later) 
in several respects, perhaps close to this form. 

4. Baikal steppe polecat, M. (P.) ev. michnoi Kastschenko, 
1910 (syn. sibiricus, lineiventer, tuvinicus, dauricus). 

Dimensions very large. Winter fur very long, somewhat coarse, 
with long shaggy guard hairs and sparse underfur. Length of guard 
hairs 64-68 mm, in posterior part of back, individual ones even 
longer (up to 80 mm). Tail densely furred with coarse hairs, its 
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 
guard hairs creates sharply bicolored fur. Dark areas between the 
fore- and hind limbs are aften united by a narrow dark band ex- 
tending along the middle of the abdomen. Summer fur (Trans- 
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 130-185 mm, females, 124-173 mm; length 
of hind foot of males is 54-80 mm, females, 48-72 mm; ear length 
of males is 24.5-26.5 mm, females, 23.5-26.0 mm. 

Condylobasal length of male skull is 64.6-82.2 mm, females, 
63.0-76.7 mm; zygomatic width of males is 37.3-58.9 mm, fe- 
males, 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; females, 16.6-19.4 mm; postorbital width of males is 12.0- 
773 17.0 mm, females, 12.0-18.0 mm (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 espe- 
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 places, the described form may show several devia- 
tions in color and measurements; however, really convincing data 
about this are lacking. It is possible that some populations have 
transitional characters with other forms. Thus, perhaps that is the 
way things stand in the northern parts of Trans-Baikaliya, where it 
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.) ev. larvata Hodgson*, 1849. 

Dimensions very large. Color light — underfur almost white, 
guard hairs black in color, but quite sparse. Red color tones absent. 

*In original text, misspelled "Hodgon" — Sci. Ed. 



1164 



There are no data about body dimensions, Condylobasal length 
of male skull is 75 mm, and apparently even more, and female is 
up to 70 mm and may be more (material of Z[oological] M[useum 
of] M[oscow] U[niversity]). 

In Alai valley. 

Outside the USSR — in Tibet, Himalayas, Kashmir and, appar- 
ently, montane parts of Kashgaria. 

This form is very little known both as regards its characteris- 
tics and also its distribution. It belongs to the group of large races 
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 revi- 
sion of Central Asian polecats. It is possible that all polecats from 
Tibet through the Pamir-Alai, Tien Shan, Altai and Mongolian 
Republic to Pri-Baikaliya themselves {michnoi) represent one form. 

6. Amur steppe polecat, M. (P.) ev. amurensis Ognev, 1930. 

Dimensions moderate, apparently less than in preceding form. 
Pelage is short, of equal length over the entire skin very dense and 
soft, 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 its whole length. Dark tip 
very short — about 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 100- 
140 mm; length of hind foot is 49-58 mm; length of ear is 22-28 
mm. 

Condylobasal length of male skull (4) is 64.1-66.4 mm; 
zygomatic width is 23.5-24.4 mm; interorbital width is 19.3-20.1 
mm. Weight of males (5) is 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 spe- 
cies, it separates itself most sharply from all others. This applies 
both to general color tone of fur, also to difference in tail color; 



1165 

and to very characteristic structure of winter fur. Although as 
pointed out, some characteristics of this form are revealed in gen- 
774 era! fur tone of animals from Trans-Baikaliya, the Amur polecat 
differentiates itself from all other races or groups of races. It may 
be considered a representative of its own group of races. In this, 
it is, apparently the only form. 



Steppe polecats living outside borders of our country have 
been very poorly studied, and their independence needs confirma- 
tion for both their morphology and distribution. Usually, the fol- 
lowing 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 is a common, locally an abundant 
predator; however, its populations are unevenly distributed, which 
is associated with the disposition of suitable habitats and their 
qualities, mainly with respect to abundance of principal foods. 
The polecat is particularly numerous where relatively large steppe 
rodents are abundant — ground squirrels and hamsters, etc. — and 
pikas. 

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 sec- 
tions, 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 USSR: in the pre-revolutionary 
years, the average annual harvest was about 900 thousand indi- 
viduals. 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 terri- 
tory — 4.1, for Kazakhstan — 1.56, and for former Nizhne-Volzhsk 
territory — 0.87. 

Habitat. The steppe polecat everywhere prefers open habitats — 
level and foothill stepppes, fallow fields, pastures, slopes of mead- 
ows 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 in which the steppe polecat lives 
will correspond to that inhabited by one or another of those ro- 
dents species constituting its food objects, and therfore it is dif- 
ficult to describe — it is variable" (Sviridenko, 1935). The steppe 
polecat ascends mountains to an altitude of 1860 m in the northern 
Caucasus, and to heights of 2000-2200 m in the mountains of 
Kazakhstan. 

Food. The steppe polecat is a typical carnivorous animal. The 
polecat cannot assimilate plant food and it is absent in the diet 
(Zverev, 1931). Invertebrates are encountered rarely in summer 
foods, and only in individual sections of the range. 
775 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 
marmots and they only feed on the young and injured animals and 
on dead bodies. 

The best conditions for polecats occur in those places densely 
inhabited by ground squirrels. In these cases it feeds on them al- 
most 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 



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 num- 
bers, density, and in the character of daily and seasonal activity, 
absence of stable settlement, etc. 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. 

Fish, domestic fowl and carrion become of some significance 
for the polecat along the shores of rivers and lakes, or with insuf- 
ficient regular foods. Birds are of secondary importance in the 
food of the steppe polecat. Among harvested birds, it attacks gray 
partridge [Perdix sp.] and willow grouse [Lagopus lagopus], but 
rarely grouse. 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. undulatus] and 
marmots, in forests — on hamsters, small voles and wood mice in sandy 
regions — on great gerbils [Rhombomys opimus], etc. 

Even primary foods (occurrence not less than 14-16%) cannot 
be considered significantly stable in its winter diet. Over eight 
winter seasons in Volga-Kama 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 in only two 
seasons out of eight (each). This is associated with fluctuations in 
abundance of these groups of food resources in nature (Grigor'ev 
and Teplov, 1939). 

In captivity, adult polecats ate 55-150 gm of food daily, on 
average — 90.6 gm, and when fed only on beef they ate an average 
of 165.8 gm. Young polecat at age seven months ate an average of 
145.5 gm daily and at the age of 2.5 months — 155 gm. When fed 
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 Nizhne- Northern Western Trans- 

territory Volga Kazakhstan Siberia Baikaliya 

territory 



Winter Summer Winter Winter Winter Winter Summer 

Marmot — _ _ _ _ 0.26 13.2 

Ground squirrel 3.86 47.0 — 76.3 1.49 1.04 13.2 

Hamster 5.69 47.1 — — 23.3 — — 

Pika — _ _ — — 76.2 15.4 

Water vole 10.66 _ _ _ 10.07 — — 

Great jerboa 0.92 — — 6.8 2.45 0.26 2.2 

Hamsters — _ _ 5.2 9.8 4.32 4.4 

Mouse-like 59.4 52.9 12.5 6.8 31.3 13.6 4.4 

rodents 

Mole-voles 0.36 — — 2.6 — — — 

[Ellobius] 

Birds 7.53 4.9 7.4 — 1.7 3.0 2.2 

Harvested birds 3.27 — 0.8 — 1.16 — — 

Common 2.0 — 0.8 — 0.15 3.0 — 

partridge 

Domestic fowl 0.73 — 5.8 — — — — 

Small birds 0.36 — 0.8 — 0.45 — — 

Frogs 4.4 _ _ _ 0.07 — — 

Fish 5.5 — 29.1 — 0.22 — — 

Carrion — — 23.7 — 0.97 — — 



Insects — 


— 


— 


— 


— 


— 


12.2 


Amount of data 550 


103 


118 


38 


1348 


766 


45 



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 independ- 
ent 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 young ground squirrels; 2) 13 ground squirrels; 3) 
5 ground squirrels; 4) 5 hamsters (Zverev, 1931). In Volga-Kama 
territory (Grigor'ev and Teplov, 1939), stores contained: 1) 4 
vipers; 2) one hamster and 4 water voles; 3) 3 water voles and 2 
gray voles. Probably, these stores that occur are not always 
utilized. 

Home range. The steppe polecats do not belong among those 
animals with a sharply-defined home range. In the warm time of 
the year, especially in places rich in ground squirrels, old animals 
hold to a relatively settled mode of 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 m from their burrows. Young polecats do not systematically 
attack all inhabited burrows of ground squirrels successively; they 
are more active, most often overnighting in the burrow of ground 
squirrel 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. 

With available food in abundance and shelters, the range of 
daily activity in summer is small. In winter, steppe polecat moves 
12-18 km per day, often returning to the same burrow from which 
it set out to hunt. The area of daily activity was determined in one 
case as 3 km^ (Zverev, 1931). 

In winter, with worse conditions for capturing food, greater 
depth of snow cover, the animals move to areas more favorable 
to them. In such case, significant aggregations are 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 by steppe polecats, their dimen- 
sions, depth, 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 many outlets (from 
3 to 15-20); moreover, alongside the nest chamber is a second 
chamber — for storing fresh food. Lining of the nest chamber does 
not occur. If one is 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, 
the diameter of passages in hard soil is 9-12 cm, in female bur- 
rows — 7-8 cm (in ground squirrels, 5-6 cm, rarely 7 cm). In soft 
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 shel- 
ters in places where existing shelters are absent. 

Daily activity and behavior. The steppe polecat has no sharply- 
defined rhythm of daily activity. It is mainly a crepuscular-noctur- 
nal 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. 
Apparently, the short summer night is not sufficient for capturing 
food, especially for females nourishing large broods. Hunting by 
entire broods was observed during the day more than once. 

The influence of weather on the activity of the steppe polecat 
has been professed, but is not strong. In winter, it is decreased 
at — 30°C and lower. Daytime activity in summer is mainly ob- 
served in cloudy, not very hot weather and in winter, mainly dur- 
ing snowstorms or in foggy, cloudy weather. According to data 
from laboratory investigations (Kalabukhov, 1943), the rhythm of 
nocturnal activity in the steppe polecat is less well-defined than in 
the forest polecats. They are considerably more active in daylight 
hours. From 22 to 55% of all activity proceeds during the period 
from 0600 to 1800 hours (36.5% on the average against 17.9% in 
the black polecat). Females are more active at night than males. 

The steppe polecat is an active, clever, brave and curious ani- 
mal. It swims excellently and climbs well, but rarely use these 
abilities. They dig out the burrows of ground squirrels very rap- 
idly. It catches its food in burrows and on the surface by waiting 
and hiding. While hunting ground squirrels, the polecat does not 
behave in a specific manner: after taking a certain direction, it 



1171 

makes a great rush, often passing over inhabited burrows. While 
hunting, the polecat searches clefts, burrows and every object that 
attracts it, even if it is not experiencing hunger. In winter, when 
hunting mouse-like rodents, it makes numerous "plunges" under 
the snow (up to 18-20 in one hunt). Deep snow makes such 
means of catching food of little productivity, and the polecat is obliged 
to change its habitat to a less 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. 

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 among mustelines is nearly the only 
778 species forming, with [prey] 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. 

Seasonal migrations and transgressions. Quickly destroying 
its 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 
not live in one place for a long. time, especially the young. It 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 relation- 
ship 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 to 
unfavorable conditions for obtaining food, causing such movements 
(deep snow, see above). In autumn, polecats 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 
sexual cycle (Kler, 1941). The period of complete sexual dormancy 
lasts 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. Exter- 
nal signs of estrus appeared on 12-13 March and developed for 2- 
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 is repeated as is also the case after death 
of a brood. In the latter case, it may be repeated for three succes- 
sive pregnancies. Usually, it 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 Zoo, the body length of newborns is 5-6 cm and 
body weight is 10 gm. These figures fluctuate depending on the 
number of young in the litter (Krumina, 1934). 



1173 

After three days, a thin white underfur appears on the body 
surface. On the 9th day, the body length doubles and body weight 
increases six-fold (33 gm). At the 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 ap- 
pears. At that time, the young polecats weigh 70-72 gm. Teeth 
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, and 
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 
mm 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 
of 60 days — with adults. They live in the family burrow for 2-2.5 
months. Dispersal begins from July, or later depending on time of 
delivery*. Young polecats living independently are already ob- 
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 per year: in spring and in au- 
tumn. 

Enemies, diseases, parasites, mortality, competitors, and popu- 
lation dynamics. The steppe polecat has many enemies: wolf, red 
fox, corsac fox, imperial eagle and snowy owl. However, their role 
in the 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 in contact with the 
majority of species of small carnivores, except for solongoi and 
weasel, only at the ecological borders of its distribution within the 
range. 

Among infectious diseases, the polecat is weakly susceptible 
to rodent [sylvatic] plague, tularemia and viral disease — 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 
scheme of the track. Voronezh district, 8 January 1952. Sketch by A.N. Formozov, 
about 2/3 's natural size. 



by more than 11 flea species, partly transmitted to polecats from 
rodents on which they feed. 

There are no data on mortality. 

Population fluctuations in steppe polecats are clearly evident. 
They are induced by food deficiency, epizootics and the death of 
780 both adult 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 
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 is well noted, but the range of fluctua- 
tions is less than that in other carnivorous animals in this region. In 
1939-1948, these fluctuations were 11 times higher due to the very 
high rise in their number in 1939, which was associated with a rare 
abundance of the Daurian pika [Ochotona daurica] and groups of 
gregarious vole [Microtus socialis]*. 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 long- 
legged buzzard [Buteo hemilasius] and almost coincided with changes 
in the numbers of Daurian pikas and narrow-skulled voles. 

Field characteristics. It is not difficult to distinguish the steppe 
polecat in open habitats. 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; 
the length of its leap is 40-65 cm (Formozov, 1930). In moving, 
it 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 mouse- 
like rodents, it can destroy about 1500 specimens in the winter 
alone (Sviridenko, 1935). In regions with weakly developed agri- 
culture, the steppe polecat is very useful, destroying natural foci 

*The Russian word is "stadnykh" — Sci. Ed. 

*°In an active season, one ground squirrel eats about 6 kg food, 25% of 
which consists of grains. 



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 its skins has occupied 
seventh place; in Kazakhstan and several other places, the steppe 
belt polecat occupies one of the first places. Its durable beautiful 
skin was always in demand in the national and international mar- 
kets. However, in past years, from 1926-1929 to 1956-1959, the 
catch of steppe polecat noticeably decreased. Along the Cis-Cau- 
casus, the catch decreased 4 times; in Bashkiria — 6 times, in Mid- 
dle Povolzh'e — 10 times, in the republics of Middle Asia — 14 times, 
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 squir- 
rels, hamsters 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. 

Harvesting this animal is simple and profitable; it is mainly 
781 taken with jaw traps placed near the inhabited burrows. In the 
northern Caucasus, up to 100-150 were caught in one season us- 
ing 10-16 jaw traps (Sviridenko, 1935), in Nizhnyi Volga — up to 
60 polecats by one hunter. 

The best method for maintaining the number of steppe pole- 
cats at the desired level, which is usually determined by the op- 
posing interests of fur trade and agriculture, is periodic prohibition 
of its harvest for 2-3 years. The prohibition must include not less 
than a whole district or territory, otherwise, it will not be effec- 
tive. In districts of field-protecting forest belts, it 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 
Gueldensta edt. 



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. Protuberances, crests etc. of skull well devel- 
oped — occipital well-defined, sagittal relatively weak, but quite 
pronounced, anteriorly bifurcating and giving off branches to post- 
orbital processes. Brain case relatively small, not elongated, quite 
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 — line of 
lateral borders of this part of skull begin to come together at an 
angle and then diverge again. Least width in region of postorbital 
constriction less than interorbital width. On the whole, this region 
is relatively short. Skull noticeably flattened. Line of upper profile 
of skull relatively straight. It is 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. 

Auditory bullae moderately swollen, somewhat angular (irregu- 
lar triangular) in outline, only 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 relatively weak, paroccipital (exoccipital) 
processes — well developed. Bony palate wide. 

3 13 1 
Dental formula I-C-P — M — = 34. Teeth are strong — rela- 
tively a little stronger than in genus Mustela. Canines, especially 
upper, are longer. Main cusp of last lower premolar (third tooth 
behind canine — in front of carnassial) has no additional cusplets 
on inner side. Lower carnassial tooth bears small additional cusplet 
on inner side of main cusp. Upper carnassial tooth is strong — 
width of its cutting part approximately equal to half of its length. 
Upper molar (last in the series) has inner and outer sides of equal 
782 length, or inner side is slightly more massive — last tooth is not 
constricted appearing as two blades (inner and outer), or constric- 
tion is ill-defined. 



1178 



Trunk thin and elongated, limbs short, tail relatively long (about half 
length of body plus head). Head small, blunt; ears large 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. 




782 Fig. 276. Reconstructed range of genus Vormela Bias., and reconstructed range of 
marbled polecat, Vormela peregusna Gueld. V.G. Heptner. 



1179 

A terrestrial animal, associated with open, arid, level areas and 
with burrows of mass settlement of rodent species (ground squir- 
rels, gerbils). A specialized predator, feeding mainly on rodents. 

Range of the genus relatively small and occupies southeastern 
Europe, parts of Asia Minor, Middle Asia, Kazakhstan, and north- 
ern parts of Central Asia (for details see following description of 
species range). 
783 Genus is monotypic. 

Genus Vormela is very well and strictly characterized, and of 
its independence no one raises any doubts. It long ago — evidently 
even in the Pliocene — constituted a group, with its origin probably 
in Asia Minor (fossils are known from the Pleistocene in Syria, 
Palestine and the Caucasus). In its 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 its main features — its entirely peculiar color — is set to one 
side, it is most similar to the steppe polecat. Speaking essentially 
of skull structure, actual fundamental differences are only in struc- 
ture 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 all 
of its 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- 
tures (fusion of pterygoid processes with 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 
libyca auct.) and is usually placed between them in the system. 
Sometimes, Vormela is considered more closely related to Poecilictis 
than to Putorius (Mensel, 1881). The whole question of generic 
groups closely related to Mustela requires special study. 

In composition of fauna of the USSR there is one species: 
marbled polecat, Vormela peregusna Gueldenstaedt, 1770. j 



1180 

Found in steppes and deserts of southern European part of 
USSR, Causasus, Middle Asia, and Kazakhstan. 

A fur-bearing species, also of significance as a regulator of 
numbers of several rodents, in particular reservoirs of harmful 
infections (V.H.). 

MARBLED POLECAT, PEREVYAZKA' 

Vormela peregusna Giildenstaedt, 1770 

1770. Mustela peregusna. Giildenstaedt. Novi Comm. Acad. Sc. 
Imp. Petrop., 14, 1, p. 441. Steppes along lower Don. 

1771. Mustela sarmatica Pallas. Reise durch versch. Prov. d. Russ. 
Reiches. 1, p. 175, 453. Between Syzran' and Medveditsa 
river. 

1910. Vormela koshevnikovi. Satunin. Zool. Anz., 36, p. 59. 

Ashkhabad. 
1910. Vormela tedshcenica. Satunin. Zool. Anz., 36, p. 60. 

Tedzhensk oasis, Turkmeniya. 
784 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 i pril. stran, 3, p. 70. Starogladovsk Station on Terek 
[river]. 

1976. Vormela peregusna euxina. Pocock. Proc. Zool. Soc. Lon- 
don, p. 718. Malsosi, Dobrudzha, Romania. 

1936. Vormela peregusna syriaca. Pocock. Ibidem, p. 720. 
Tiberias* Lake, Syria. 



•Perevyazka — is now an artificial, bookish name, though in the 18th cen- 
tury, it was apparently in 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, 
the name "polecat-perevozchik" or simply "perevozchik" is used. This name is 
without meaning. It may itself represent a distortion of one of the old Ukrainian 
folk names "perevyznik" (Pallas, 1811). 

*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. Inst. AN 

SSSR, 7, p. 129. Semirech'e, former Kopal'sk со. 
1948. Vormela peregusna obscura. Stroganov. Ibidem, p. 131. 

Vakhsh valley, Tadzhikistan (V.H.). 



Only species in its genus. 



Diagnosis 



Description 



In general appearance, the marbled polecat is very similar to pole- 
cats. As in them, it often moves in leaps, usually arching the back. 
With this, however, several of its ecological particularities are 
noticeably different from polecats: instead of a sharp chirp, the 
voice is threatening — a peculiar growl or grumble, and adopts a 
special defensive pose, etc. (see below). 

Head small and narrow, muzzle obviously blunt, ears relatively 
very large — 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. 

Winter fur quite short and not very close-fitting, with rela- 
tively weakly developed underfur and dense guard hairs. Fur, not 
counting very long hairs on tail, is even, and sharp contrast be- 
tween underfur and protruding guard hairs, as in polecats, is ab- 
sent 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 summer 
fur relatively small, and 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 com- 
bination 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 
type of color which, within the family Mustelidae, is 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 is not only complicated, but 
entirely variable individually and, to a lesser degree, geographi- 
cally. In main features it may be represented by the following 
scheme (after A. A. 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 
the tail, including lower part of sides and legs, shining black color 
with light brownish tints. 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 color very clear and, contrasting, consisting of black and 
786 pure white parts. Upper and lower lips, back to corners of mouth 
and chin — "peristomal ring" — pure white. Quite wide black stripe 
passes through eyes in form of transverse band. Descending to 
sides of head and reaching corners of mouth, it unites with black 
throat area. Behind this black band, a quite broad pure white 
transverse stripe runs between eyes and ears — "frontal band". On 
reaching lateral head surface of the head, it slopes somewhat 
posteriorly, passing below ear and ending on sides 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 
narrow middle one that 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 form- 
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 its great variability. For additions, see below, and section on "Geographic 
variation". 



1184 

nape corresponding as it were, to frontal band. Together with 
lateral stripes (in these cases, the middle one is poorly defined), it 
forms an angular bracket-like figure opening posteriorly — "collar". 
Color of neck stripes white — somewhat yellowish in posterior part, 
and main tone of this part of neck brown or rusty-brown. 

Whole dorsal side from shoulder region to base of tail or 
almost base, occupied by variegated saddle-patch. It spreads 
ventrally and laterally, especially in posterior half of body, where 
it 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 actual saddle-patch itself represents a large area of 
basically 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 vari- 
able 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, more or less rounded spots of brown 
color are scattered over it. They also sometimes display a tendency 
towards fusion into longitudinal, oblique, and even transverse 
groups. The anterior part of the saddle-patch is commonly more 
densely covered with spots; in its posterior portion they are smaller, 
more sparsely scattered, and the yellow field of the saddle-patch is 
paler, on the whole, more clearly defined and more striking to the eye. 

Both described types of saddle-patch themselves represent 
negatives of one another — 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, 

*In the Russian original, "analnykh (poroshitsevykh)"; both words appear to 
have the same meaning — Sci. Ed. 



1185 

the remains of the main dark background of saddle-patches of the 
first type. On the whole, saddle-patch color is to the highest degree 
variable, both in general features and details. From the color type 
in which all basic elements of the saddle-patch are expressed, are 
787 derived the origins of all forms; one side, as far as a large, light, 
area with insignificant dark spotting, and on the other hand — up to 
a dark 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. 

Base of tail is rusty-brownish or brown in color. Its middle — 
the longest part — is a pale-straw 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 
at base region darker than upper, usually bright blackish-brown 
and on remaining part, as on upper side. 

Vibrissae and hairs covering soles of feet dark-brown, foot 
pads not covered with hairs in winter. Claws light, 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, some- 
times 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 charac- 
ter; 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 structure of the os penis is in the main similar to majority 
of species in genus Mustela in particular, to the polecats. Its base 
is laterally compressed, and the distal third is bent upward hook- 
like. The dorsal surface of the basal half carries a rib-shaped el- 
evation, and the end is flattened. The very tip itself forms a rounded 
denticle slightly bent to the right. There is no groove on the lower 
surface of the basal part, but terminally it is well-developed. 

Dimensions of the marbled polecat are not subject to great 
individual variation; in particular, especially large individuals have 
not been noted. 

Body length of males is 269-352 mm, of females, 290-345 
mm; tail length of males is 1 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 
Z[oological] M[useum of] M[oscow] U[niversity]). 
789 Condylobasal length of male skull is 50.6-62.0 mm, of fe- 
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, of females, 13.1-16.0 mm; mastoid width of males is 26.1- 
32.6 mm, of females, 27.2-30.2 mm. 

Os penis length is 36.9-39.2 mm (3 individuals; Ognev, 1935). 
General weight is 370-715 gm (V.H.). 



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 (reconstructed) constitutes a considerable part of the 
range of the species and itself represents the north of its western 
half. 

In the west, the northern 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 
region (Voronezh preserve) and thence, in an imprecise way pro- 
ceeds to the Volga — to Syzran' or a little south of Syzran'. Beyond 
the Volga the border, apparently, rises north of 52° [N. lat.] and 
directs itself to Orenburg on the Ural river. 

South of this line, the range in the west extended, apparently, 
to the foot of the Carpathian mountains but in these mountains, the 
marbled polecat was evidently not present. It was also not re- 
corded in the Trans-Carpathians. The range probably extended into 
Romania in the extreme lower Danube. Apparently, the marbled 
polecat existed all 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 ex- 
tends to the shore of the Caspian Sea. In the Caucasus, the marbled 
polecat is distributed over the entire Cis-Caucasus steppes from 
the mouth of the Kuban' to Makhachkala, reaching the foot of the 
mountains (Nal'chik, Vladikavkaz). In places, it goes up into the 
foothills somewhat (Psebai on the Belaya — a tributary of Kuban'), 
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 
Derbent, and farther south, to Baku. 



1189 

In the Trans-Caucasus, the marbled polecat inhabits the plains 
and foothills in the Kura and Araks basins, passing along the Kura 
to Tbilisi. In Armenia, it is met with along the valley of the Araks 
and the montane steppe region and in semideserts north of it, 
proceeding into the western part of the country (to the 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. 
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 the Zaisan depres- 
sion, and here exits beyond the borders of our country to Dzhungaria 
[China]. 

To the south of the above-mentioned line, in the Asiatic part 
of the country, the range occupies the whole of the Middle Asian 
plains. The southern border either goes beyond the borders of our 
country, or passes along the foot or foothills of the southern moun- 
791 tain systems. 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 south- 
ern border skirts the Tien-Shan system; apparently, the Fergana 
Valley together with Karatau are included in the range, and it 
skirts 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. 

To the west of the Amu-Dar'ya, the range of the marbled polecat 
exits beyond state borders everywhere. It is, however, absent in the 
Kopet-Dag, although in the lower western parts of the range, it 
penetrates into the mountains, notably along the foothills and val- 
leys. 

The assumption about the inhabitation of the marbled polecat 
in the Trans-Urals to 54° 50' N. lat. (Kundrava southeast of Chel- 
yabinsk — not 55° 30' as written by Ognev, 1935) is a mistake and 
is, apparently, based on an imported skin. The affirmation of its 
existence around Baikal, whence a separate form was even de- 
scribed, and in "Siberia" (Pocock, 1936) is not well-founded — this 
species is 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 in Orlovsk district, 
the marbled polecat still existed, although it was rare, but in 1925, 
it had already completely disappeared. At that time or somewhat 
later, the 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 
the Sea of Azov to Osipenko city or a little westward — to the 
meridian of Melitopol.^ 
792 In the interfluve between the Dnepr and Volga, the northern 

border, begins near Voronezh, descends somewhat southward, and 
then, passing slightly north of 50° N. 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 — these are Poltava (perhaps even a more or 
less permanent occurrence), and Pugachev in Trans-Volzh'e. 

On the whole, the border of the range for the last 100-200 
years receded very strongly — from north to south (not to mention 
the right 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 its occurrence also in Nikolaevsk district (Vladimirsk 
region); however, this place was not shown in the 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 lies in the 
ploughing of the virgin steppes with which it is associated, and in 
the complete agricultural utilization of the steppe and forest-steppe, 
and in particular, in the disappearance of ground squirrels. The 
area of their normal and more or less continuous distribution be- 
gins east of the Don (at Kamensk, 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 — "a fugitive of culture" — by another carnivore of 
its lineage ecologically analogous in many ways, but revealing a 
tendency towards being a "companion of culture". They secure 
for themselves subsequent prosperity where the marbled polecat 
does not persist. No competitive relationships between these spe- 
cies was observed in sufficiently extensive areas of sympatry. 

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 

In Europe the range includes Dobrudzh in Romania (in remaining 
part of 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 

"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^, Afghani- 
stan, Baluchistan, Iran, Iraq, Syria, Palestine and Asia Minor (the 
details of its distribution here are unknown)*. Probably, the 
range goes on to Kashgaria, but positive information about this is 
absent (V.H.). 

794 Geographic Variation 

Geographic variation of the marbled polecat is still poorly studied. 
In all, about 12 geographical races are known, of which more than 
half belong to our territory. This number is, apparently, exagger- 
ated, for there are not more than three real subspecies in our coun- 
try. The fundamental characteristics of all forms are based on 
coloration. At the same time, as shown above coloration of the 
marbled polecat is 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, commonly based on limited material, 
was presented by several authors, especially older ones (for exam- 
ple, K.A. Satunin) but also recent, is 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 one author (Stroganov, 1948), 
in the nominal form the white band on the forehead is broken, 
while all other authors describe it as continuous; in another case, 
the distribution of the form negans was found to be disjunct, and 
between the two parts of it lies the range of another form (Novikov, 
1956), etc. 

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 ed. Harrison 
Zoological Museum. 



1195 




793 Fig. 281. Some previous and present borders of range of marbled polecat, Vormela 
peregusna Giild, in the European part of USSR. V.G. Heptner: 

1 — Reconstructed northern and western borders of range (western more or less 
approximate); 2 — The recent (at beginning of 60' s) northern and western borders; 
3 — Separate points of occurrence in last decade; 4 — Separate distant transgressions in 
last decade (Poltava, Pugachev); 5 — Some peripheral places of occurrence in begin- 
ning of 20th century (before 1914). Map depicts formation of the break in "rounded 
pontoon" ring of original range. 



1196 

adjacent ones. It was ascertained that the main feature of geo- 
graphic variation of color lies not in detailed color difference, but 
first 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 differ- 
ences in size between races have not been confirmed, dimensions 
of the separate forms are not given. 

1. South Russian marbled polecat, V. p. 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 
distinct; 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. Bounda- 
ries with respect to the next form unknown. 

Outside the USSR — in Romania and the Balkans. 

The systematic position of the marbled polecat of the Trans- 
Caucasus has not been studied in the manner required. They have 
been related (Ognev, 1935) to the population inhabiting the east- 
ern Cis-Caucasus С intermedia') on the basis of very little mate- 
rial. It is possible that they belong to syriaca — a form which in 
general is very little known. 

2. Transcaspian marbled polecat, V. p. 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 
anteriorly by a broad transverse white stripe (forming well-defined 
"collar"). Color of lateral occipital stripes usually yellowish. Scapu- 
lar stripes very distinct; of thigh, weakly defined and bearing dark 



1197 

spots. White frontal band not divided medially and has no connec- 
tion with peristomal ring. Abdominal spots are absent or rarely 
occur, weakly defined and few (1-3). Claws short — the longest 
less than 10 mm. 

In the western part of Middle Asia — eastwards to at least Amu- 
Dar'ya and region of right tributaries of Pyandzh and upper Amu- 
Dar'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 a well-defined 
race, close to the nominal form. It is very similar to the latter in 
dorsal color, but dark tone often even darker and therefore fur 
color appears more contrasting. Except for this indistinct charac- 
ter, 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 
diagnostic significance. The author's suggestion concerning its small 
dimensions not confirmed (Chernyshev, 1956). Although the 
author considers this form "one of the most differentiated sub- 
species" (Stroganov, 1948), there is insufficient basis to separate 
it from the Transcaspian V. p. koshevnikovi. Moreover, its range 
(valleys of Vakhsh and Pyandzh) is perhaps very small. Such small 
ranges are not characteristic of races of Palearctic carnivores. 

The forms peregusna and koshevnikovi as a pair of closely 
related dark forms, contrast sharply with the very light pallidior 
and, apparently, to its pair member — negans. 

The presence of dark races which shows no essential charac- 
ters of "degradation" of color in Turkmenia and southern 
Tadzhikistan and, apparently, in Afghanistan — countries with clearly 
defined desert climate and in general, desert conditions — 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 






Ш< 




795 Fig. 282. 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 

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 Kara-kum (Uch-Adzhi; Z[oological] M[useum of] M[oscow] 
U[niversity]). The very light color of the Semirech'e and Central 
Asiatic animals clearly contrast with this form. There are, there- 
fore, two groups of race of marbled polecat — the dark European, 
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 analo- 
gous features known in some other carnivores (weasels, badgers, 
bears). 

The name adopted here for the Transcaspian form nas an in- 
disputable priority over alpherakyi* Birula, under which this form 
is usually known, although both of them appeared in the same 
year. 

3. Semirech'e marbled polecat, V. p. pallidior Stroganov, 1948 
(the name negans was applied to this form). 

797 General color light. 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 abdo- 
men and are usually larger. Claws quite long — reaching 13 mm. on 
middle digits of fore foot. 

In the region to east of the Karatau range, Semirech'e, Cis- 
Altai steppes and montane steppes of southern Altai (Chuisk). 

Outside the USSR — in Dzhungariya, and Mongolian Republic, 
apparently except in southeastern or eastern parts. 

A well-distinguished form, obviously contrasting with both 
previously described forms. It is possibly the lightest form of the 
species, or in any case of members of the group of light races (see 
above). Based on its color type, it 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. 

Despite the obviousness of characters of the Semirech'e popu- 
lation in the sense of comparison to several other races and distri- 
butions — this, of the many forms, is the least known and unclear. 
The Semirech'e marbled polecat was previously (Ognev, 1935) 
considered as transitional from the Transcaspian V. p. koshevnikovi 
Calpherakyr) to the Central Asiatic (Ordos) V. p. negans or was 
allocated directly to this race. In any event, its differences from the 
Transcaspian form were emphasized, which is completely correct. 

A 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), is not understandable. In general, the race inhabiting 
the area 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 
developed "collar"; i.e., an eastern type. However, as is known, 
the marbled polecat is absent not only in the vicinity of Baikal (if 
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 form ornata is not in any way possible 
and this name is best excluded in the category of nomen delendum. 



Beyond the borders of the USSR, the following forms are 
indicated (not counting euxina, see above, and ornata): V. p. negans 
Miller, 1910 — Ordos, Inner Mongolia, Shaanxi and some other 
parts of Inner Asia, not accurately clarified, and possibly also eastern 
or southeastern parts of the Mongolian Republic; V. p. syriaca 
Pocock, 1936 — Syria and adjacent territories (Palestine?) eastward 
to western Iraq. 
798 Note. V. p. chinensis Stroganov, 1962 described from a speci- 
men labeled "China" and earlier assigned to negans (Ognev, 1935) 
does not deserve separate status. The nature of area inhabited (and 
origin of the type) is set forth as "basin of the lower course of 
Hwang Ho". None of the arguments in use leads to such an inter- 
pretation of the label, not to mention that in the lower course of 
Hwang Ho if that part of the river below the great northern bend 
can be so considered, the marbled polecat is entirely, apparently, 
absent. 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 

Population. Almost everywhere the marbled polecat is encoun- 
tered in the European 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 Cis- 
Caucasian steppes, the marbled polecat was most common in the 
eastern part of the steppes (Kizlyarsk steppe) where it is drier. 
There it is most often met with at the present time (V.G. Heptner). 
The marbled polecat is now common in Kazakhstan, in the 
Muyunkum sands, southern Pribalkhash'e, and along the Syr-Dar'ya 
(Sludskii, 1953). The marbled polecat is most common in 
Turkmenia, where its distribution is closely connected with the 
great gerbil [Rhombomys opimus] and it is encountered formerly in 
great numbers, in the flat part of the country it was the most 
common small carnivore. Its population significantly exceeds the 
weasel population (V.G. Heptner). 

Habitat. The marbled polecat is an animal of open unwooded 
expanses — dry steppes, semideserts and deserts. It is met with 
chiefly in places rich in large desert and steppe rodents — gerbils, 
ground squirrels and in part jerboas, etc. The ploughing of virgin 
steppes and fallow lands pushes out the marbled polecat and re- 
duces its range. It is encountered in the southern grass steppes of 
the chernozem zone, does not avoid thickets of blackthorn, buck- 
thorn and other shrubs in steppe shelterbelts. It infrequently occu- 
pies afforested river valleys and the borders of forest massifs of 
799 the forest-steppe. It is encountered occasionally among gardens, 
melon fields or even in settlements; but in dry hilly steppes, it is 
rare. In Middle Asia, it mainly inhabits localities of desert and 
semidesert character, but it does not avoid river valleys and sec- 
tions cut by irrigation ditches and sometimes overgrown with a 
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 scat- 
tered 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 al- 
ways 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 is absent. 

Burrows and shelters. The marbled polecat for the most part 
inhabits burrows of various steppe rodents, only widening and 
deepening them. In Turkmenia, and generally in Middle Asia, the 
marbled polecat lives permanently in colonies (towns) of great 
gerbils, not only because of the close trophic connection with this 
rodent but also in connection with its 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 colony of great gerbil, Rhombomys opiinus. Badkhyz 

preserve, southern Turkmenia. May 1962. Photograph by A. A. Sludskii. 

ground squirrels, but this is an exception. The marbled polecat 
does not make its own burrows as a rule (V.G. Heptner). 

Daily activity and behavior. The marbled polecat is mainly 

800 active in twilight and early morning. It spends the day in burrows 
(Sludskii, 1953). In Turkmenia in winter time, spring and autumn, 
when the heat is still not too great, daytime activity occurs in the 
marbled polecat or morning hunt is sustained longer — when sun is 
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 mar- 
bled polecat rarely catches them; mainly it catches them in 
burrows (V.G. Heptner). 



1205 






rt 



^fi^^^" 



«f 









^) 



'*<» >. 






^?'^-'Й> f 



'^^- N 






»^ 

800 Fig. 286. Tracks of marbled polecat on wet sand in a gallop and scheme of leaps. Foot 
prints are placed in fours — hind 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. 

A 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 up over its 
back (like a squirrel). At the same time, it rises up high on its legs, 
jerks up its head and bares its teeth. In contrast to the "chirring" 
of polecats, ermine and weasels, it gives a peculiar growl like a 
dog. The frightening effect is strengthened by the bright, vari- 
egated black-and-white fur. When running, the marbled polecat 
expands its tail, fluffing it out. 

Seasonal migrations and transgressions. Data are lacking. 

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 

to young were caught. In the northern Caucasus, parturition 
occurs 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 
4.3. Thus, the marbled polecat is less fecund than the steppe 
polecat. 

Males, it seems, participate in raising the young. Spermatogen- 
esis begins to be extinguished in April. 

Growth, development and molt. Information is absent. 
801 Enemies, diseases, parasites, mortality, competitors, and popu- 

lation dynamics. This carnivore has no dangerous enemies. The 
steppe polecat may be named as one of the most fundamental 
competitors influencing the distribution and numbers of the mar- 
bled polecat. Other steppe carnivores hardly exhibit a noticeable 
influence as competitors. 

The marbled polecat is almost unsusceptible to plague, and 
more susceptible to tularemia. 

Human changes in the virgin steppes and their ploughing lead 
to the displacement of the marbled polecat and to a sharp decrease 
in their numbers. Utilization is not large and has no effect at all on 
the number of the marbled polecat (P.Yu.). 

Practical Significance 

The importance of the marbled polecat as a fur-bearing animal 
is negligible. In the USSR, slightly more than one thousand skins 
are taken annually. Its fundamental significance is as a predator 
which destroys steppe rodents, vectors of dangerous infections. 
Its role in destroying rodents harmful to agriculture is not great 
since the marbled polecat is rare in agricultural regions, and it 
avoids cultivated landscapes. On the whole, it is a useful animal 
(P.Yu.). 



1207 
Genus of Honeybadger or Ratel* 
Genus Mellivora Storr, 1780 

1780. Mellivora. Storr. Prodr. Meth. Mamm., p. 34. tab. A. Viverra 
ratel 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 brain- 
case. 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 swol- 
len. 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 mo- 
lars); upper first molar relatively very small and strongly elongated 
in transverse direction. On the whole, dental system is quite simi- 
lar in its main features (structure and relative size of teeth, etc.) to 
that of species of the subfamily of true martens, Mustelinae, espe- 
cially polecats. The essential difference is only in that the second 
lower molar is, as a rule, absent. 

3 13 1 
Dental formula I-C-P-M- = 32. ** 
3 13 1 

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 lysie barsuk are literally "honey- 
eater" and "bald badger" respectively — Sci. Ed. 

'The generic synonyms, since they are not used in the Russian literature, are not 
given. 

3^1 3 1 

**In Russian original, erroneously given as I — C— "P — M — = 32 — Sci. Ed, 



1208 



An omnivorous predator, mainly feeding on small vertebrate 

and invertebrate animals. Ecologically, it is an entirely flexible 

802 form, encountered chiefly in different types of flatlands, mainly open, 

to sandy deserts inclusively, but it avoids continuous moist tropical 

forests; 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 



1> 






%V^. 



, Moscow' 



eod 





•^=<5.., 




Fig. 287. Range of genus of honeybadgers, Mellivora Storr. and species range of 
ratel or honeybadger, Mellivora capensis Schreb. Schematic. V.G. Heptner. 



1209 

Within the boundary of the genus, many species were de- 
scribed, especially at the end of the last century and beginning of 
the current one, both from Africa and also from Asia. At the 
present time, it 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 con- 
sidered in doubt. However, the attempts made to establish its sys- 
tematic 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 
was assigned to the 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 
!03 Mellivora with Melinae. At the same time, the similarity between 
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, as was in part mentioned above (see also 
beyond), according to skull, both in main features and many de- 
tails of dentition, the genus reveals greatest similarity to species of 
the subfamily of martens, true Mustelinae, and is strongly differ- 
entiated 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 sepa- 
ration 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 wolver- 
ines, Gulo. If the wolverine is actually a specialized deviant 
gigantic marten, the honeybadger may be considered a huge 



1210 

aberrant polecat, including also in this designation the above-men- 
tioned (рЛ179) 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 remain- 
ing Mustelinae, the significant distance of the genus from Meles 
and the subfamily Melinae in general points to the systematic position 
of the skunk (Vonyuchka*) group, subfamily Mephitinae. They are 
clearly considered to occupy an intermediate position between 
"Mellivorinae" (honeybadger) and Melinae (badgers). However, it 
is indubitable that, according to dentition for instance, skunks are 
sharply differentiated from the honeybadger and all 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 sub- 
family of skunks is restricted in its distribution to the New World 
only, but in the past (upper Miocene-upper Pliocene) it was also 
found in the Old — in 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 rela- 
tionships 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, 
1776. 

The forms of the genus are, apparently, practically indistin- 
guishable. 

In the USSR, distributed in the western part of Middle Asia, 
and associated with deserts and desert mountains. 

It has no practical significance (V.H.). 

^According to their dentition which may consist of 28, 30 and 36 teeth in 
different forms, they are even 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. pi. 125. Also: 1777, 3, 

pp. 450, 588. Cape of Good Hope. 
1792. Ursus indicus. Kerr. Animal Kingdom, p. 188. India. 
1851. Mellivora ratel. Horsfield. Cat. Mam. E. Ind. Co., p. 120. 

India. 
1920. Mellivora wilsoni. Cheesman. Journ. Bombay Nat. Hist. Soc, 

27, p. 335. Ram-Khormuz on the Iran-Iraq boundary. 

Diagnosis 

Only species of the genus. 

Description 

In general proportions, appearance, and behavior, the honeybadger 
is entirely unique and, in spite of the prevalent opinion, does not 
resemble the badger. It more quickly reminds one of a bear-cub on 
the one hand, and on the other, it has much in common with the 
polecat. It often moves in a sort of creep, extending its body, 
bending its 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 rela- 
tively 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 
hairs of upper foot. 

'"Medoed" [Honey-eater] is an artificial name, representing a translation of the 
Latin generic name. Another name exists, 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. 



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 — Kashan 
interfluve, southern 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. 

In winter fur, entire lower body — belly, throat, chin, lower 
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 
as well as anterior part of muzzle including eye region — is a shiny 
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 reach- 
ing 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 
or flask-shaped form with a narrow "extension" on the tail. 

The general tone of this area is white, but in details of color, 
the dorsal saddle is somewhat variable. Hairs on the head and 
806 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 
distal half and with a wide dark base. These bases more or less 
(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, es- 
pecially with a strong admixture of dark in the saddle patch, is 
separated in the form of a fuzzy lightening on the general back- 
ground, 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 
same color as the middle of saddle patch. Sometimes in its poste- 
rior part, a lighter narrow border is noticed. In different individu- 
als, the dark tone is visible through the white, sometimes more and 
sometimes less, and the general color is lighter or darker but these 
differences are, apparently, small. 

Sexual differences in color are absent, and age differences are 
undescribed. Among the exotypic variations, malanism is known 
(Africa). The geographic variation in color, if present, is very weak. 

Skull relatively large and quite heavy, of entirely unique struc- 
ture'*. The upper profile of the skull presents itself as a quite even, 
arched line having its 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, al- 
though wide, is short; its length is approximately two times shorter 
than the braincase. 

The postorbital constriction is long and very narrow — its width 
is always less than that of the interorbital area, usually consider- 
ably so (by 5-10 mm). The braincase is large — elongated and wide, 
broadening posteriorly and swollen in the parietal region. The 
occipital crest is strongly developed, and the sagittal is relatively 
weak (much weaker than in the badger) even in the very old ani- 
mals. Zygomatic arches are strong but not very massive, in the 
medial portion arching upwards very strongly. The lateral lines of 
the interorbital region are parallel or almost parallel, the supraorbital 
processes have wide bases, but are short. 

"The skull of the honeybadger is often compared to the skull of the badger as the 
most similar. This 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. 



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 is greater than the zygomatic, or at 
least equal to it. 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 auditory bullae very large, swollen, angular-oval and 
strongly diverging anteriorly — their anterior walls adhere closely 
808 and partially fuse with inner parts (those directed towards sagittal 
plane of skull) of postarticular processes. Hard palate short and 
wide; its interior width between molars 2.5 times less (or about 
that) than distance from posterior median sphenopalatine notch to 
alveoli of middle incisors. Portion of palate lying behind toothrow 
relatively short 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. 
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 consider- 
ably 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 
Mustelinae, and also in badgers and, in part, in otters, there is no 
tendency towards reduction or loss of this tooth. 

Sexual differences in skull structure are absent or not notice- 
able, and no age variation has been described. 

The OS penis has, in terminal quarter, a зЬаф bend upwards — tip 
located nearly at right angle to axis of proximal half. Tip itself not 
evenly divided — one "horn" is larger than the other (Pocock, 1941). 
In general, structure is typical for martens and polecats (subfamily 
Mustelinae) and differs strongly from that characteristic of badgers. 

The body length of males (2) is 680-750 mm, of females (1), 
680 mm; tail length of males is 180-205 mm, of females, 172 mm; 
length of hind leg in males is 120-125 mm. 

'It 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 
females (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 
82.8-86.4 mm, of females, 80.7 mm; interorbital width of males 
(4) is 31.7-33.8 mm, of females (2), 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 mm; length of upper toothrow (without incisors) 
in males (4) is 38.5-40.2 mm, of females (2), 30.8-37.2 mm. 

Weight of male is 12 kg (September) and 16 kg (October), of 
female, 9.1 kg.^ 

There is sexual dimorphism in body and skull measurements — 
the females are smaller than the males — but they are relatively 
insignificant. Difference in overall weight is more obvious (V.H.). 

Systematic Position 

Only species in the genus. 

Geographic Distribution 

Africa, a large part of Near Asia, India, western Middle Asia. The 
range of the ratel is one of the most extensive of the species ranges 
in the family. 

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 [range] and nearby 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. 



1219 

The western border on the north is formed by the Caspian 
coast, to a little north of 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 depres- 
sion, and through Deukal village (Deukaskenkala, Onkhauz and 
Doudyr), generally at the latitude of Kunya-Urgench, and some- 
what to the west of it reaches the Amu-Dar'ya delta. In the delta 
itself and in its contiguous cultivated lands, in particular the Khiva 
oasis, the honeybadger is not, apparently, encountered. From the 
Kunya-Urgench district, the border turns to the southeast, becom- 
ing the eastern border of the range and, apparently passing around 
Khiva oasis, and going farther along the Amu-Dar'ya, reaches the 
state border. In the middle course of the river, the ratel was 
recorded at Darganat (tugai) and in the upper course, it was 
recorded, as noted above, at Kalach near Kerka. 

The honeybadger has not been recorded on the right bank of 
Amu-Dar'ya and is, apparently, absent there. The Amu-Dar'ya 
constitutes an insurmountable barrier for this species dispersing 
from the south — 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 soils. Within the 
boundaries of the described region, the ratel is known, for 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 others'. 

Beyond that, in the beginning of the 90s of the previous cen- 
tury, when the first information on the occurrence of honeybadgers 
within the borders of our country appeared (Bikhner and Zarudnyi, 
1892; Varentsov, 1894) there was no information at all about it for 
a long time, and the prevalent opinion was that it appeared occa- 
sionally among us, or in 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 
that it occurred at furriers in these places starting from 1924, 
although in 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 — "very rare", 
"exclusively southern Turkmenia"). 

Investigations in the 50' s and the very beginning of the 60' s 
and, to a lesser extent, in the 40' s, showed that the ratel is not at 
all 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 
result of wide zoological investigations which have unfolded 
in 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 

''The honeybadger is found in a great number of places along the above- 
mentioned 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 lie at a distance ranging from 350 km (at the 56° 
E. long.) to 650 km (along longitude 59° E. long.) from the pre- 
viously known ones (lO's and 20' s) and the more eastern parts 
even farther — up to 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 
its northern and northeastern parts. Despite being a large eurytopic 
species, sandy deserts are for it, 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 
in Egypt and reached the Mediterranean Sea in this part of Africa. 
However, the northern border of the range in Africa is quite in- 
definite. Judging by features of the species described from the 
Trans-Caspian part of the range, in the east, it apparently passes 
generally keeping to the 20° N. lat. along the southern edge and in 
the southern parts of the Sahara, and in the west, it is, apparently, 
more northerly — along the hills of the Ennedi desert, Tibesti, Adrar- 
Iforos and others. In any event, its presence was established on the 
Air plateau (Asben, 1800 m above see level). Thence, the border 
of the range passes in an unclear way through the western parts of 
the Sahara to southern Morocco (Ifni, Sus and the Valley of Um- 
er-Ryabii, a little south of Casablanca). In the Sahara, the ratel 
only avoided probably, its northeastern parts, notably the Libyan 
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 in Iran as very limited and clearly 
incomplete, believing that it may, however, exist in other places. According to this 
author, honeybadger inhabits only a small territory in the extreme southwestern corner 
of the country (in Khuzistan) between the head of the Persian Gulf and the southern 

contd. on next page 



1222 

Hindu Kush), India, from the former Northwest Frontier Prov- 
inces to Bengal inclusive (absent in Assam and Burma), northward 
to the foot of the Himalayas and the southern zone of Nepal, in the 
south to the sea coast — to the southern extremity of the peninsula. 
It is, apparently, absent in the Malabar coast. In Ceylon, it is ab- 
sent. In the described reconstructed range it has apparently not 
undergone essential changes, except in Egypt (V.H.). 

Geographic Variation 

With so significant a range, the ratel undoubtedly displays known 
geographic variation; however, it is not, apparently, great. Thus, 
the differences 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 is only one form, 

apparently, not differing in any substantive way from the Indian. 

Indian honeybadger, M. с 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 to exist in the territory of 
Hindustan — indica and inaurita. They differ from each other only 
in that in the second (eastern) the fur is somewhat denser, and 
hairs are found on the heel. It is assumed that this form may also 
be found in the provinces contiguous to Afghanistan (Pocock, 1941) 

contd. from previous page 

half of the border with Iraq. On the other hand, its range was described as a narrow 
belt along our borders from Tedzhen to the Gorgan (Astrabad) Gulf on the Caspian 
Sea. Judging by the distribution of the species in the USSR and in neighboring 
countries and by its biological characteristics, one may allow that it 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-e- 
Lut). 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 more severe con- 
ditions (winter) than those under which the Indian form lives, it is 
possible that the fur of the former is denser than that of true indica, 
i.e. it is like inaurita. However, the differences themselves be- 
tween these forms are doubtful: our museums have no materials of 
these races and therefore the name given is retained for our ani- 
mals. The relationship to the form wilsoni from southwestern Iran 
is perhaps unclear, but judging from everything, this form, de- 
scribed from individual characteristics of one specimen, does not 
have substantial truth. 



The following forms are usually described from parts of the 
range lying outside the USSR. Independence of a series of these 
forms, including Palearctic ones, is doubtful. 1) M. с inaurita 
Hodgson, 1836 — Nepal and parts of the range east of it^ ; 2) M.c. 
wilsoni Cheesman, 1920 — southwestern Iran, Iraq (?); 3) M. c. pumilio 
Pocock, 1946 — Hadramawt, southern Arabia; 4) M. с leuconota 
Sclater, 1867 — West Africa northwards to southern Morocco; 5) M. 
с signata Pocock, 1909 — Sierra Leone; 6) M. с consica Thomas et 
Wroughton 1907 — Lake Chad; 7) M. с buchanani Thomas, 1925 — 
Air plateau (Asben), Sahara; 8) M. c. abyssinica Hollister, 1910 — 
Ethiopia; 9) M.c. broekmani Wroughton, 1920 — Somalia; 10) M. с 
cottoni Lydekker, 1906 — Ituri region, Congo; 11) M. с makwelli 
Thomas, 1923 — Kenya; 12) M. с sagulata Hollister, 1910 — Tangan- 
yika; 13) M. с vemaj/ Roberts, 1932 Bechuanaland; 14) M. с capensis 
Schreber, 1776 — South and Southwest Africa. 

The actual number of forms of honeybadgers is, evidently, 
smaller. This is clear at least 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 its range, 
the honeybadger may be called a eurycious species. It is met with 
in sands of the Karakum; in sands and foothill plains of the Kopet- 
Dag range; in montane canyons of the latter; among steppes and 
hills of Badkhyz; in marsh tit* plains covered with saxual, Russian 
thistle [Salsola] and sagebrush; among deep ravines of the south- 
813 ern precipice of Ust'yurt [plateau] with stunted vegetation of shrubs, 
sagebrush, and succulent saltwort; in cultivated river valleys among 
the bai** landscape; and in dense riparian tugai thickets. 

Food. Insufficiently studied. In India, it feeds on small ro- 
dents, birds, reptiles including cobra, as well as grasshoppers, miner 
bees, wasps and their larvae and honey. In India, they sometimes 
attack poultry-yards. Its bias towards 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, Ust'yurt, Kazakhly village. 
Photograph by V.P. Kostin. 

*The Russian word {pukhlyakoba) means marsh tit, but its significance as 
a descriptor of landscape is unclear — Sci. Ed. 

**bai is a Turkic word referring to a wealthy landholder in Middle Asia; 
i.e., river-valley land — Sci. Ed. 



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 it also eats hedgehogs; 
a case of eating the carcass of a lamb was observed (Kostin, 
Sapozhenkov, Gorelov, Zhernovoi and Svyatoi, 1963). Among rep- 
tiles, feeding on the desert monitor was recorded in Badkhyz (Sukhinin 
and Shcherbina, 1955 and others). It very often feeds on Horsfield's 
tortise which it digs out even in winter. Small lizards and round-head 
lizards [Phrynoce-phalus] also occur in its food. At the southern edge 
of Ust'yurt (V.P. Kostin), up to 30% reticulate scales of 
Phrynocephalus were found in feces. Among insects, grasshoppers 
were found, in particular the saxual humpbacked grasshoppers, and 
darkling beetle (Sapozhenkov, Kostin, Zhernovoi and Svyatoi, 1963). 

Home range. Information is lacking. On the edge of the Ust'yurt, 
tracks of honeybadger were traced for a distance not less than 2.5 
km; it follows that, the home range is quite large (V.P. Kostin). 

Burrows and shelters. The honeybadger lives in burrows dug 
by itself. It digs very well, and is 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 
rhythm of daily activity. As a rule, it leads a crepuscular-nocturnal 
515 way of life (Sapozhenkov et al.; 1963; V.P. Kostin), but is some- 
times 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. A male caught on 10 October, had large testes. 
In Ashkhabad Zoo, a female caught on 6 May delivered on 



1226 




X 



^A 



^ 




814 Fig. 293. Habitat of 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. 




!14 Fig. 294. Hilly landscape in Badkhyz preserve, southern Turkmenia. Habitat of 
ratel or honeybadger. May, 1961. Photograph by Yu.K. Gorelov. 



1227 




\-0 



-fO 



-10 



CM 



k^ 






Fig. 295. Tracks of honeybadger. Sketch by V.P. Kostin. 



1228 

31 May; she gave birth to one cub. In the first days of May, a 
lactating female 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 popu- 
lation dynamics. Data are absent. 
816 Field characteristics. The color of the honeybadger masks it 

very well in solonchaks. Moving in an uneven gallop, it is very 
reminiscent of the general form of the wolverine. Prints of its 
tracks are wider and shorter than the badger. Its feces have a 
sausage-like form, 4-7 cm long and 12 mm diameter. Becoming 
angry, it raises its tail upward when walking (V.P. Kostin). 

In addition to the gallop, by which the honeybadger can pro- 
ceed for a prolonged time, it also walks and even half-crawls, 
sprawling on the ground. The voice of the honeybadger resembles 
a hoarse "khrya-ya-ya-ya". At the time of rut, the male makes a 
loud grunting sound (Sapozhenkov et al., 1963). The honeybadger 
climbs trees well. It gives off a frightening smell and exceptional 
movement of the skin surface (P.Yu.). 

Practical Significance 

Due to its small numbers and because it inhabits little-populated 
places, it has almost no practical significance. It is not hunted espe- 
cially and is for the most part captured accidentally. Honeybadger are 
sometimes caught alive for zoos. For this, the animal's tracks are 
followed and they are dug out from burrows (Sapozhenkov et al., 
1963). (P.Yu.). 

Subfamily of Badgers 

Subfamilia MELINAE Burmeister, 1850 

Genus of Badgers 

Genus Meles Brisson, 1762 

1762. Meles. Brisson. Regn. Anim., p. 13. Ursus meles Linnaeus, 
1758. 



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; 
tail short. Limbs not shortened, massively plantigrade with long 
claws of digging type; webbing between digits absent. Ear pinnae 
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. Be- 
tween the anal opening and. root of tail there is a glandular "pocket", 
into which an odorous secretion exudes. Anal glands well-devel- 
oped. Teats 3 pairs — 2 abdominal, 1 inguinal. 

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 is considerably 
less than zygomatic. Auditory bullae of moderate size and not 
swollen. Auditory tube not covered by mastoid process and opens 
independently in front of it. 
Я7 Dentition is a specific type, fundamentally differing from that 

characteristic of Mustelinae. It represents a combination of teeth of 
a sectorial type with those having clearly defined structural fea- 
tures of a tuberculo-grinding (crushing) tooth. The structure of the 
dentition is one of the most characteristic features of the sub- 
family. It is more similar to the dentition of the subfamily of otter 
(Lutrinae). Upper carnassial tooth not largest in toothrow — it is 
short and triangular in form. Upper molar is largest, considerably 
larger than carnassial, flat, tuberculate and of grinding type. Lower 
carnassial tooth elongated. All cheek teeth, except upper molar, 
are of more or less distinctly sectorial type. 

3 14 1 
Complete dental formula is I-C-P — M — = 38. However, 

individual first premolars or all of them are frequently lost and 



1230 

the formula changes correspondingly (see below, description of 
species). 

They are 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 capa- 
bility 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 consid- 
erable part of Siberia, Near, Central and Eastern Asia as well as 
Middle Asia. 

In the current century, several species were included in the 
genus other than the European M. meles — Cretan M. arcalus, Asi- 
atic M. leptorhynchus, Tibetan M. leucurus and Japanese M. 
anakuma. For our fauna, two species — M. meles and M. leptorhyn- 
chus — 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 denti- 
tion (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, 
the genera Taxidea (American badger), Arctonyx (Indian "sand" 
badger), Helictis (including Melogale:* "ferret" badgers of south- 
eastern Asia) and Mydaus (teledu — Malay archipelago). All of them 
are sharply distinguished from each other (Helictis is sometimes 
separated in an independent subfamily) and it is difficult to estab- 
lish their systematic relationship. 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 somewhat inferior to 
Meles